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animal life

A stunning pictorial essay . . . that serves as a key to the animal kingdom.

N e w Yo r k T i m e s B o o k R e v i e w




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First American Edition, 2008

First paperback edition, 2011
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Editor in Chief Dr. Charlotte Uhlenbroek

Richard Beatty

Chris Mattison

Dr. Sean Rands




Dr. Frances Dipper

Dr. George C. McGavin

Dr. Graham Scott

Fishes, Defence

Invertebrates, Predation, Scavenging

Dr. Kim Dennis-Bryan

Dr. Sanjida OConnell

Feeding, Feeding on Plants, Omnivores, Feeding




Professor Tim Halliday

Dr. Douglas Palmer

Amphibians, Sex and Reproduction,

Reproducing without a Mate,
Finding a Mate, Sexual Rivalry,


Dr. Elizabeth White

Steve Parker

Vision, Birth and Development,

Raising Young

Skeletons and Muscles, Movement,

Body Coverings, Body Systems

John Woodward

Rob Hume

Dr. Katie Parsons


American Museum
of Natural History

Dr. Juliet Clutton-Brock

Dr. Christopher J. Raxworthy


Chief consultant

Dr. Frances Dipper

Dr. George F. Barrowclough


Professor Tim Halliday

Dr. Randall T. Schuh

Dr. Mark E. Siddall

Rob Hume

Dr. John S. Sparks

Chris Mattison

Dr. George C. McGavin




Senses, Living Space, Life Histories


Dr. Charlotte Uhlenbroek

Dr. Robert S. Voss

Animal data
The behavior proles in this book contain summary
information about the animals being described.
This information usually covers a species, but in some
cases it refers to a group, such as family or genus.
The symbols used in the summary panel are as follows:
a In most cases (and unless otherwise stated), this
refers to the length of the adult animal, with the
following dimensions given for different groups:
Mammals: head and body
Birds: tip of bill to tip of tail
Reptiles, Amphibians, and Fishes: head and body,
including tail
Invertebrates: head and body, including tail but not
including antennae
f Occurrence, including both habitat and geographical

c A brief description of the animals main physical
dCross-references to other behavioral proles about
the same species or group

Foreword by Charlotte Uhlenbroek




What is an animal?


Animal history




Animal groups
26 Invertebrates
30 Arthropod groups
32 Arthropods
36 Vertebrate groups
38 Fish groups
40 Fishes



Animals bodies allow them

to maintain a stable shape,
move around, and sense and
react to their surroundings.




Animals have lived on Earth

for more than a billion
years, and have evolved a
staggering array of forms.


82 Water skeletons
83 Horny skeletons
84 Chalky skeletons
86 Bony skeletons

92 Climbing, leaping,
and jumping
94 Burrowing, slithering,
and sliding
96 Flying and gliding
98 Swimming

Body coverings
102 Skin
104 Scales

46 Amphibians

106 Feathers

50 Reptile groups

107 Fur, hair, and bristles

52 Reptiles
58 Birds

90 Walking and running

44 Amphibian groups

56 Bird groups

Skeletons and muscles


Body systems
110 Breathing

68 Mammal groups

112 Circulation

70 Mammals

113 Digestion
114 Fluids and temperature
116 Brains, nerves,
and hormones

120 Touch and vibration
122 Taste and smell
124 Vision
126 Hearing
128 Echolocation
130 Electricity and magnetism




An animals behavior
encompasses all the things
it does, from competing for
food to forming societies.





380 Life stories

138 Home ranges

and territories

396 Raising young

148 Migration

408 Play and learning

162 Animal architects







182 Feeding on plants

194 Omnivores
202 Predation
250 Scavenging
260 Feeding partnerships


276 Weapons and threats

444 Pheromones
and smell

296 Camouage and


448 Visual signals

312 Group defense

456 Sound
462 Touch, vibration,





328 Reproducing without

a mate
332 Finding a mate
340 Sexual rivalry
352 Courtship
366 Mating







I am thrilled to have worked on this book. It is unique,
so far as I am aware, in exploring the richness and
diversity of animal behavior across the animal kingdom.
The essentials of animal life are relatively simple
nding food, self-defense, reproduction, and
dispersalbut the ways in which animals achieve
these things are extraordinarily complex and varied.
From amazing hunting strategies and extraordinary
defenses to stunning mating rituals and complex relationships, the
wealth of different behaviors reects the diversity of animals and the
huge range of physical and social environments in which they live.
Animals populate every corner of the globe from the bottom of the sea
to mountain peaks and from the Equator to the poles; they may live in
groups of thousands or meet others of their kind just once in a lifetime.
An animals behavior depends to a large extent on its form and
evolutionary history, and the rst two parts of this book provide a clear
and comprehensive description of the major groups of animals and their
evolution, anatomy, and senses. The main part of this book then looks
at animals in action. It is divided into eight chapters that look at different
aspects of animals lives, although these are not always mutually
exclusive. It doesnt attempt to provide a complete catalog of all animal
behaviorthat would run to many volumes and is expanding all the
time. Nevertheless, it is an extensive overview of our current knowledge
of animal behavior, including typical and unusual behaviors in different
animal groups, classic studies, and new discoveries.
It is an exciting time to be writing about animal behavior. Advances
in neurobiology, anatomy, and physiology have revolutionized our
understanding of behavioral mechanisms; the rapidly expanding eld
of ecology is revealing the extraordinary web of relationships between
species; and the maturation of long-term studies of social behavior is
providing intimate views of animal societies. Many discoveries are blurring
the boundaries between human and animal behavior, such as the use
of symbolic signals to communicate, intense sociality, and problem
solving. Meanwhile, new technology has taken us into realms utterly
alien to our experience such as the use of polarized light and magnetism
to navigate, electricity to hunt, or minute vibrations to communicate.
This book reects the extraordinary body of work carried out by
biologists who have devoted years, often whole lifetimes, to the patient
observation of animals in the eld and in the lab and to designing
ingenious experiments to learn why animals behave in the ways they
do. Some species have been very well studied and we know an
extraordinary amount of detail about their behavior; others are barely
known and as yet we only have tantalizing glimpses into their lives.
Written by a team of respected biologists and illustrated with fantastic
photography that jumps off the page at you, this book really puts the
life into wildlife. The great biologist E.O. Wilson said, Once you know
an animals behavior you know its essence. I hope that this book
helps capture the essence of animal life on this planet and makes us
ever more determined and equipped to protect it.



What is an animal ?
There are at least two million living species of animals on Earth. They range from microscopic
worms to huge whales, but they all share a few key characteristics which, when combined,
distinguish them from all other forms of life. These dening characteristics include both
physiological features and the behavior that makes animals so intriguing.
Life on Earth is traditionally divided
into ve kingdoms. Two of these,
consisting mostly of bacteria and
protists, are mainly microscopic, so
we are rarely aware of them. The
other three are more familiarthe
fungi, plants, and animals. The
essential differences between fungi
and plants may not be obvious at
rst sight, but most animals are easy
to recognize by the way they move
and react to their environment.
Some aquatic animals move very
little, and may look a lot like plants,
but animals and plants function in
very different ways.



Mammals 9%
Birds 19%
Reptiles 13.5%
Amphibians 10%
Fishes 48.5%
Arthropods 60.5%
Cnidarians 0.5%
Worms 3%
Echinoderms 0.5%
Mollusks 5%
Other 30.5%

Scientists believe that there may be more than ten
million species of animals, although fewer than
two million have been described scientifically. Of
these, less than 3 percent are vertebrates, such as
mammals, birds, and fishes. The arthropods make
up by far the largest group of animals on Earth.

Energy and food

Plants, and many bacteria and protists, use the energy of sunlight to make
food. They combine carbon dioxide with water to form a sugar called
glucosea simple carbohydratein a process known as photosynthesis.
The glucose stores energy, which plants use to fuel their growth. They can
also convert glucose into other carbohydrates, such as the tough cellulose
that reinforces their structure. As they absorb the water they need, plants
also acquire dissolved chemicals
such as nitrates and phosphates,
which they turn into proteins.
Animals cannot make their own
food from simple chemicals. A
few, such as reef corals, live in
partnership with organisms that can
produce food like this, but most get
their nourishment by eating plants or
other living things. They digest the
living tissue to break it down into
simpler ingredients, such as glucose,
and use these to fuel and build their
Gerenuks are superbly adapted to reach the most
own bodies. Their need to nd food
succulent young shoots on trees. Standing on their
is one reason why animals have
hind legs, they can stretch their long, muscular
evolved mobility and senses.
necks in among the branches.

Unique embryos
Some simple creatures, such as jellysh, sea anemones, and corals,
can multiply asexually by growing buds that turn into new animals. Many
insects and other invertebrates, such as water eas and some aphids,
can develop from unfertilized eggsa process known as parthenogenesis.
In both cases the young are clones of their parent.
However, most animals produce single-celled eggs, each fertilized by
the sperm cell of another individual to create a zygote that contains the
DNA of both parents. This single fertilized cell develops into a ball of cells
called a blastula, which is unique to animals. Through a process of cell
division, the blastula becomes a multicellular embryo (see p.379). Having
inherited a mixture of genes from each parent, the embryos produced in
this way are genetically different (except in the case of identical twins).
Combined with gene mutation and natural selection (see pp.1415), this
genetic mixing creates the variation that has enabled animals to evolve into
such a dazzling variety of species.

Animals are built up from many microscopic cells which,
unlike those of plants or fungi, do not have rigid cell
walls. In all animals, except sponges, the cells are
organized into different types of living tissue such
as muscles and nerves. These tissues may form
specialized organs such as the heart, brain, and
lungs. The body plan usually becomes xed early
in life, but the bodies of some animals, such
as butteries, undergo a radical rebuild, called
metamorphosis (see p.386), when they become adults.

All animals obtain nutrition from other living organisms
or from the remains of dead organisms. Parasitic
worms that live inside other animals absorb simple
nutrients through their skins. Other animals have
ways of ingesting food into gut cavities, which are
specialized for digesting it and turning it into useful
nutrients such as glucose. Some aquatic animals
simply lter food particles from the water, but most
animals have well-dened, mobile mouths that they
can use to seize and even chew their food.

Gas exchange
All animals need oxygen to turn carbohydrate food into
energy, a process that releases carbon dioxide. This
is the reverse of photosynthesis, which produces
oxygen, so animals are oxygen consumers, not
producers. They exchange gases through thin-walled
gills in water (as in shes), through their moist skins
(as in amphibians), through branched tube systems
(insects), or through lungs. Many types of animal have
bloodstreams of some description that carry these gases
around the body, along with food, such as blood sugar.

Sensory systems
Nearly all animals, except sponges, have networks of
nerve cells that respond to external stimuli. Touch
a simple animal like a sea anemone (a cnidarian),
and it will twitch. More complex animals have
sense organs that react to light, sound, pressure,
scent, taste, and even electricity. They also have
brains that can memorize and recognize the stimuli,
enabling many animals to learn by experience. Most
of these sensory organs are concentrated at the head
of a typical animal, near its mouth.

The most distinctive feature of animals is their mobility.
Some aquatic animals, such as mussels, spend their
adult lives attached to rocks and may not move
visibly, but they do pump water through their bodies.
Mussels also open and close their shells as the tide
rises and falls. Most other animals are able to slither,
crawl, swim, walk, run, and even y. Combined with
their senses and memories, this enables them to seek
out food, escape enemies, and nd breeding partners.
In other words, they display behavior.

What is evolution?


These three moths are members
of the same species. The
variations originally occurred as
a result of genetic mutations.
Natural selection led to the
darkest moth becoming
more abundant, since
industrial pollution
made them blend
in better to the
blackened trees
that the moths
rest on during
the day.

Evolution is the process

of change in the inherited
characteristics of populations
of animals over time. The
crux of the theory is that
all life today has evolved
and diverged from simple
ancestors that lived in the
oceans over 3 billion years
ago. This means that all
animals are related to each
other. Accumulated evidence
from biology, genetics, and
fossils supports evolution as the
unifying theory that directs our
understanding of life and its history.


Charles Darwins denition of evolution was descent with modication,

a term he used in his book The Origin of Species to describe how
successive generations of a species adapt to their changing
environments over time to eventually form entirely new species.






This is often described as the survival
of the fittest. This species of butterfly
varies in color and reproduces in large
numbers, not all of which survive, due
to limited resources in the environment.
Predators eat more of the purples, which
are less camouflaged than the yellows.
The surviving yellows produce more of
their own kind. With continued predator
preference over time, the yellow
butterflies become dominant.

Macroevolution and microevolution










The large-scale pattern of change is known as

macroevolution. Macroevolution refers to large-scale
changes, such as the evolution of limbed vertebrates
from those with ns. Other examples of macroevolution
are the emergence of the shelled egg, freeing some
land-living, four-limbed animals from dependence upon
water for reproduction and the divergence of egg-laying
reptiles into other major groups, including turtles and
crocodiles. This occurs as a result of small-scale
descent with modication, known as microevolution.
For example, since their introduction to the United
States, house sparrow populations in the north and
south of the continent have developed differences, with
the northern variant becoming biggerprobably as an
adaptation to the colder climate.


To the left is a cladograma diagram showing
a group of organisms descended from a common
ancestor. The animal groups in the diagram are all
related to the first vertebrate, which appeared around
540 million years ago. The branching patterns occur
as a result of divergent evolution. These branching
patterns form a family tree(or phylogeny).

Randomly occurring change in the
genetic makeup of a population
over time is known as genetic drift.
For example, a forest fire wipes
out most of the purple butterflies
in the population. The next
generation contains the genes
of the lucky survivors, not
necessarily the fitter ones.
Despite being unlikely, it is
even possible for a series
of chance events to lead
to the total loss of the
purple population.
Gene flow (or
migration) results
from the movement
of genes from one
population to another.
For instance, genes are
carried from one population
and introduced to another
by the migration of an adult
organism. In this example, a







The genetic uniqueness of
individual humans and all other
sexually reproducing organisms
results from the reshuffling of
parental genes. Offspring are
not genetically identical to each
other (except identical twins),
nor to one or other parent, but
show various combinations of
their parents genes. New gene
combinations, and hence
genetic variation, is introduced
into a population through the
mechanism of sex.



Every life form is made up of a specic
series of molecules, and the order of
molecules is contained in a chemical
code. This code is extremely complex
and is encased in spiral-shaped
molecules of deoxyribonucleic acid
(DNA). Chemicals, known as bases,
link each molecule of DNA. There are
four different kindsadenine, cytosine,
guanine, and thymine. They are always
linked up in pairsadenine always
fuses with thymine, and cytosine with
guanine. The sequence of these bases
makes up the cells genetic code.
The code found in each human cell
consists of 20,00025,000 separate
instructions. Each of these instructions
is known as a gene, and each gene
is responsible for controlling particular
characteristics. For example, there
is a single gene that is responsible
for the color of eyes. Genetics is the
study of how these characteristics
are inherited and is one of the central
pillars of biology.

each DNA
forms a
known as a

the DNA
molecule forms
a spiral shape
(a double
helix), and is
linked by the 4
different bases



leaves a purple
population and
joins a yellow
population. The migrant
interbreeds with members
of the yellows and, in doing
so, introduces its purple genes
into the yellow population.

A mutation is brought about by change in the genetic
material of an organism that is subsequently inherited
by its offspring. This chance alteration can happen
through deletion or insertion of a single base in a DNA
molecule (see panel, right). Occasionally, single
mutations may produce large effects but generally,
evolutionary change is the result of many mutations.

Among other meanings, a

species can be dened as
a group of similar organisms
that can interbreed to
produce fertile offspring.
Speciation is the process
whereby new species evolve
Isolated gene pools in islands such
from a single ancestral
as the Galapagos create unique
species. This occurs for a
traits. This woodpecker finch has
number of reasons, including evolved to use tools to catch prey.
geographical isolation arising
from habitat fragmentation. If a small population is
isolated from the main group and its members can only
share genes with each other, over time they will evolve
independently to the point where, if they came back in
contact with the original group, they could not interbreed.

Divergent evolution

Over vast periods of time, repeated speciation has led

to evolutionary divergence, where new descendent
species become signicantly different from their common
ancestors. For instance, all life on land has diverged from
water-living ancestors, and all living species of mammal have
diverged from a common ancestral mammal that lived alongside
the dinosaurs. These series of changes arise as a product of
Earths dynamic environments, which vary from place to place
and from time to time.

Convergent evolution


Sometimes, different organisms will

evolve similar characteristics to adapt
to the environment that they inhabit.
For example, the similarity in body forms
of whales, seals, and penguins is the result of
similar adaptations to similar ways of life. The
common factor is the adaptation of a streamlined
body shape, with reduced limbs for a more
efcient swimming motion. In a similar way,
wings have evolved independently in birds
and bats for ight.


Some flowers rely on hummingbirds for pollination,
while some hummingbirds rely on specific flowers
for nectar. They have coevolved in terms of shape
and color to accommodate each others survival.



This chart shows the five basic mechanisms that enable
evolutionary change to happen. The butterflies in the diagram
(like the peppered moths on the far left) are all members
of the same species, so they are capable of
interbreeding. The butterflies were originally
yellow, but a chance mutation in the genetic
material of one of the yellows resulted in
the birth of a purple butterfly (as shown
in the mutation segment). Over
sufficient time, it is possible that
the purple and yellow butterfly
numbers could be even.








One third of all brachiopod and bryozoan families, and
many groups of conodonts and trilobites were rendered
extinct. Overall, around 100 families of marine
invertebrates were wiped out.

Animal history
The evolution and expansion of animal life on Earth is a
remarkable story. From microscopic beginnings billions
of years ago, animal life evolved in the protective and
supportive medium of ocean water. Fossilized remains
show us that the earliest multicellular creatures evolved
in the Ediacaran period, around 630 million years ago
(MYA). However, the fossil record is by no means
complete. We know surprisingly little about the origins of
some groups, such as sharks and rays, but with each new
fossil discovery, we move a step closer to answering the
questions that have puzzled paleontologists for years.


How this chart works

This chart separates animal life into three major
habitatssea, land, and air. The major animal
groups are individually colored. Their relative
abundance through time is represented by an
expanding and contracting band. The evolution
of each group is signified by tie lines that link

them to their ancestral group. The three

invertebrate bands are shown on a separate
scale to the vertebrates due to the difficulty in
estimating their numbers (they are thought to
account for 97 percent of all species). The size
of their strands are relative to each other.

Diversication and proliferation

Early in the Cambrian, marine invertebrates underwent a period of
expansion and diversication. Many familiar living invertebrate groups
evolved in this period, along with now extinct groups. The rst
vertebrate animals also evolved in the Cambrian. These diversied into
several sh-related groups, some of which are now extinct. After the
arthropods moved into the terrestrial environments, they proliferated
and rapidly diversied. By late Devonian times, tetrapod (four-limbed)
vertebrates had left the water and the next stage in vertebrate evolution
had begun. The airways were the last environment for animal life to
conquer, with insects rst achieving this in the Carboniferous period.
Many animal groups have members that have evolved and adapted to
life in each environment. For example, reptiles rst evolved on land,
then evolved water-living members before nally taking to the air.









Earliest terrestrial arthropod groups

(centipedes, millipedes, and others)
evolve around 450 MYA.

Extinction events
The history of animal evolution was not one of simple expansion and
diversication. As animal groups evolved and died away, changing
environments and events impacted upon their evolution, sometimes
disastrously. There were several major extinction events that reset the
evolutionary clock. The most devastating of thesethe Permian
Triassic extinctionwiped out 96 percent of all marine species.



The first complex multicellular organisms began to
appear in the Ediacaran period, around 630 MYA .
This fossil of a Mawsonites species is thought to
be an early jellyfish or a primitive worm.

Earliest jawed fishes

evolve around 450 MYA.

Evolution of coral


The Cambrian
Explosion, around 540
MYA, saw the rapid
evolution of groups of
marine invertebrates,
and the evolution of the
first vertebrates.


Primitive nautiloids
(marine cephalopods)
become abundant
around 475 MYA.

Coral reefs become

widespread in the
mid-Ordovician period,
around 470 MYA .









This event mostly affected marine organisms. Coral
reefs, brachiopods, and trilobites were all severely
reduced in number. Most terrestrial animals were
unscathed, but some early amphibians were wiped out.

Evolving from fishes, the first

tetrapods (four-legged
vertebrates) rose up out of the
water and started colonizing
the land around 370 MYA .
Acanthostega had both lungs
and gills, eight digits on each
limb, and webbed feet.


Late Carboniferous
insects begin to appear
around 320 MYA,
including roaches
and dragonflies.
Archimylacris was an
early cockroach with
folded wings.

This fossil of Westlothiana

lizziae was hailed as the
first true reptile due to its
superficial reptilelike
features, but it is now
thought to be an amniote
ancestor (340 MYA ).


Amniote ancestors are

thought to branch out
from the amphibians
around 340 MYA.


The amniotes split into two groups around

315 MYAthe synapsids (eventually leading
to living mammals) and the sauropsids
(eventually leading to living reptiles).

Early land snails evolve

in the Carbonifeous,
around 320 MYA.

Earliest known
scorpions evolve
around 418 MYA.



Arthropods continue to diversify.

Graeophonus was a true spider relative.





Acanthodians (early jawed fishes), such as

Cheiracanthus, expand early in the Devonian
period, around 410 MYA .



Colossal early Permian sharks, such as

Heliocoprion, become the dominant
predators in the seas, around 298 MYA .



Early jawless fishes are common around

405 MYA . Pterapsis had a distinctive flattened
head, enclosed by massive bony plates.

Extensive development of
Carboniferous coral reefs,
around 350 MYA .





The first true flies evolve in the mid-Triassic.

The earliest known mothArchaeolepis
mane, evolves around 190 MYA.

Pterodactylus was a
pterosaurthe only
nonavian reptiles to
develop powered flight.
Their wings were formed
of membranes of skin.

Birds evolve around

155 MYA . Archaeopteryx
had more in common
with dinosaurs than
modern birds.


Cynodonts were a group of synapsids,
which are thought to be direct ancestors
of mammals. Cynognathus was a
carnivorous doglike cynodont.

Dinosaurs evolve around

230 MYA .


Mammals continue to
diversify. Amblotherium was a
small, primitive insectivore.

Compsognathus was one of the smallest

dinosaurs to have existed. It was chicken sized,
but extremely agile. It mostly preyed on insects.

Postextinction expansion
of dinosaurs and other reptile
groups, around 200 MYA .



The first marine turtles

evolve around 215 MYA .

Bony fishes rapidly expand and diversify after

the PermianTriassic extinction. Lepidotus
was a common bottom-dwelling fish with
thickly-enamelled scales.

Large marine predators thrived

around 175 MYA . Ichthyosaurus had
strengthening bones around its
eyes, enabling it to
dive very deep.



Triassic recovery in the seas. Reefs

emerge again with new kinds of coral.



Earths most severe extinction event, affecting all animal
groups. 96 percent of marine species were wiped out,
and around 70 percent of terrestrial vertebrates.


23 percent of both marine and nonmarine families were
wiped out, including sponges, gastropods, bivalves,
cephalopods, brachiopods, insects, and vertebrates.


Cretaceous spiny lobsters,

such as Eryon, were common
in shallow waters. The family
still exists today, but they have
adapted to the ocean depths.






Flying insect
groups, including
march flies,
continue to
expand rapidly.

The earliest known bee

Melittosphex, evolves
around 100 MYA .






Rise of primitive seabirds, around 95 MYA .


Modern bats evolve, around 40 MYA .




Homo sapiens evolved 150,000 years ago.

Tyrannosaurus rex was one of the largest-ever
terrestrial carnivores, evolving around 67 MYA .






Many features of Cretaceous
period bony fishes, such as
Hoplopteryx, are shared by
their modern ancestors.


Toothed whale ancestors,

including Prosqualodon,
appear around 30 MYA .


Squalicorax was similar
to the modern tiger shark,
with triangular, flattened
teeth and finely
serrated crowns.

Stingrays are common in the

oceans. Some, like Heliobatis,
spread to freshwater rivers.







Unusually shaped
Cretaceous starfish,
such as Metopaster,
are common in the
worlds oceans.


All non-avian dinosaurs were wiped out, as well as flying
reptiles. Marine invertebrates were seriously affected.


Before 1650, the study of living organisms
was much more localized than it is today.
However, when the early explorers started
to send home vast collections of exotic
plants and animals previously unknown
to science, it soon became evident that
without some sort of ordered system, the
situation would rapidly become chaotic.

The Linnaean classification system is shown here in
its original form. To demonstrate how this system
works, the highlighted boxes trace the systematic
position of the Indian rhinoceros, from the broadest
grouping of kingdom to the narrowest grouping of
species, which comprises only Indian rhinoceroses.

Order from chaos

John Ray (16281705) was the rst person to attempt
to classify the natural world. He organized organisms
based on their form and structure, or morphology,
using lengthy names that incorporated a brief anatomical
description. But it was Carl Linnaeus who devised the
system of classication that we still use today. Like Ray,
Linnaeus made use of morphological features, but used
them to group things together rather than to describe
them. He set up formal categories on the basis of
shared morphological features, creating a hierarchy of
increasing exclusiveness that extends from kingdom to
species (see diagram, right). Over time, scientists have
expanded the system, adding levels such as domain
and cohort, and subdividing others into infra-, super-,
and sub- categories to accommodate our increasing
knowledge of different animals. Despite these revisions,
the Linnaean system has remained fundamentally the
same since its inception 250 years ago.

In formulating his hierarchical system, Linnaeus also
streamlined the names of individual organisms
species were previously referred to by a common
name or descriptive anatomical phrase. He adopted
Latin as the universal language of taxonomy, and gave
each taxon a unique two-word name, called a binomial,
by combining the genus and species names. Homo
sapiens, for example, is the scientic name for humans.
What makes the name unique is the species partall
humans carry the generic name Homo, including fossil
humans such as Homo habilis, but only modern humans
are referred to as Homo sapiens, or knowing man.
Binomial names can still be descriptive but, more
crucially, the unique name avoids confusion.

4 classes

12 classes

4 classes

28 orders

29 orders

4 orders

Tree shrews
2 families

18 families

1 family

Carl Linnaeus (17071778) was a
Swedish botanist. Often referred to
as the father of taxonomy, Linnaeus
published the first edition of the Systema
Naturae, his classification of living
organisms, in 1735. The resulting
system remains in use today.

Nature does not proceed

by leaps and bounds.

The common name robin is applied to very different
birds, but using their Latin names readily distinguishes
them. The American robin (left) is Turdus migratorius,
while the European robin (right) is Erithacus rubecula.

12 phyla

Indian rhinoceros

2 species

Sumatran rhinoceros

3 phyla

35 phyla

14 phyla

30 phyla

This is the highest level of

Linnaeuss hierarchy. Each
contains living organisms that
work in fundamentally the same
way. Initially, there were only two
kingdoms, plant and animal,
but today there are ve.

3 classes

2 classes

7 classes

17 classes

A major subdivision of the animal

kingdom, made up of classes.
Phylum Chordata, for example,
comprises all animals that possess
a precursor of the backbone,
called a notochord, at some
time during their lives.

3 orders

Cartilaginous fishes
c.10 orders

Bony fishes
46 orders

1 order

A taxonomic level made up

of orders and their respective
subgroups. Class Mammalia,
for example, comprises those
chordates that have a single
jaw bone (the dentary), fur, and
mammary glands.

18 families

Odd-toed ungulates
3 families

16 families

2 families

More exclusive are the different

orders into which a class is
subdivided. Each order contains
one or more families and their
subgroups. The Perissodactyla, for
example, are plant-eating mammals
that walk on an odd number of toes.

7 genera

4 genera

5 genera

A family is a subdivision of an order,

and it contains one or more genera
and their subgroups. Family
Rhinocerotidae, for example,
comprises those perissodactyls
(odd-toed ungulates) that have
horns on their noses.

Sumatran rhinoceros
1 species

White rhinoceros
1 species

Black rhinoceros
1 species

Aristotle (384322BCE) was the

rst to use the term genus to
group things together. It was later
adopted by Linnaeus to identify a
subdivision of a family. The genus
Rhinoceros contains the onehorned rhinoceroses.

This group comprises similar
individuals able to interbreed in
the wild. Indian rhinoceroses, for
example, breed only with one
another and not with other types of
rhinoceroses. Their species name
unicornis refers to the single horn.

The presence of vertebrae
instead of a notochord
characterizes all groups
except the hagfishes
on this cladogram.
Possession of jaws
is the character used
here to unite all
groups including and
below sharks.


A bony skeleton rather
than a cartilaginous
one separates the
sharks and rays from
ray-finned fishes.

Lobed fins
The precursors of
limbs, lobed fins are
not found in ray-finned
fishes but are present
in the other groups
in some form.





The main part of this diagram shows part of the chordate cladethe
vertebratesas it would appear in a simplified cladogram. Read from
the top downward, each new character relates to a point of divergence from
the group to the left, which lacks the evolutionary innovation in the form
displayed by the groups to the right and below. Here, the hagfishes diverge
first, followed by the lampreys, sharks and rays, and so on down
to the mammals.

A relatively new system for classifying living organisms,
which arose in the 1950s, is called phylogenetics, or
cladistics. Based on the work of German entomologist
Willi Hennig (19131976), it unites organisms in groups
called clades on the basis of morphology (form and
structure) and genetic characters. The system assumes
that a character shared by a group of organisms but
not by others indicates that they have a closer
evolutionary relationship to each other and therefore a
more recent ancestor in common. Like the Linnaean
system (see pp.2021), this method of classication is
hierarchical, but, unlike those created by Linnaeus, the
groupings are used to construct taxonomic trees using
evolutionary relationships.

Primitive and derived characters

The characters that are important in cladistic
classication are referred to as derived because they
have altered in some way from what is considered the
ancestral primitive condition. They also need to
be present in at least two groups, or taxa, to be
informative about relationships. For example, in the
rhinoceroses, the character of a hairy body is found
only in the Sumatran rhino (Dicerorhinos sumatrensis)
and therefore it tells you nothing about its relationship
with the rest of the rhino species, whereas a single
horn (see character 4 in the diagram, opposite) is
unique to both the Indian (Rhinoceros unicornis) and

Javan rhinoceroses (Rhinoceros sondaicus), suggesting

that they inherited it from an ancestor they shared in
the past. Derived characters, such as a single horn,
that are shared among taxa are referred to as
synapomorphies. Although, for simplicity, only single
characters are shown in the rhinoceros diagram
opposite, the number of characters used to create
a cladistic hierarchy, or cladogram, is usually very
largeso large that it takes a computer to analyze
the data and generate the cladogram.

Common ancestry
Cladistics assumes that the more derived characters
species have in common, the more closely related
they are to each other than to anything else. This being
the case, it also then follows that they have a more
recent ancestor in common than they do with other
taxa. This can be conrmed by examining the fossil
record for taxa or characters once a cladogram
has been generated. Brothers, for example, share more
characters with each other than they do with their
cousins because they have the same parentsparents
being their common ancestors. They also have features
in common with their cousins because they share
the same grandparentsso the whole family would
be placed within the same clade, but with the cousins
branching off earlier from the line leading to the
brothers. In the same way, rhinoceroses form a clade
within the odd-toed ungulates.


Until recently, systematics was based on
morphological characters, because
investigation of evolutionary history involves
looking at fossils in which DNA is not
preserved. Today, cladistics is being used
increasingly to examine the relationships
of living animals. For these organisms,
relationships can be established using
DNA analysis. Such work has led to major
revisions in some of the traditional
groupings. For example, whales are now
grouped with even-toed ungulates and
more specically with hippopotamuses.

Species 8



Organisms that eat other organisms for nourishment

and whose bodies are built of many cells, all of which
lack cell walls.

Animals with integrated multicellular body structure, with

cells organized in tissues. Eumetazoans include all animals
other than sponges, which lack tissues.



Species 20,000

Species 9,000


Species 17,500

Species 17




Species 1

Species 2,000

Species 12,000

Lophotrochozoans whose ancestor evolved a chalky shell,

a muscular foot, and a body gcover called a mantle.
Not all molluscs retain these.

Species 1,200


Bilaterians whose larva (young form) is of a unique type
called a trochophore. This group is dominated by molluscs
but has other important members.


Species 90

Solenogastres, Caudofoveates
Species 750

Species 20



Species 2,000

Species 4,300


Species 650



Species 300


Species 1,450


Species 350

All descendants of the rst eumetazoan to evolve a
body with a single line of symmetry, featuring a front
end, usually with a head. Bilateria includes nearly all
animals, even those that have a radial (wheel-like)
arrangement, such as starsh.

Bilateral (symmetrical) animals with a three-layered outer
covering that is moulted. Most ecdysozoans are arthropods,
but roundworms are also abundant.

Group descriptions
Major groups have a
short description of
key characteristics.

Smaller groups branch
off at the appropriate
points. The scientifc
name and number of
species is given for each.


Species 320

Species 2,000

Species 20,000

Species 150

Species 35,000


Species 100





Species 14,000

The layers of color reflect
relationships. For example,
vertebrates are part of
craniates, chordates,
deuterostomes, bilaterian,
eumetazoan, and animals.

Species 100

Species 1



Species 1,400

Species 100,000

Species 70

Bilaterians with a radial pattern of cell division in the early

embr-ude some invertebrate groups plus chordates the
phylum containing

Deuterostomes with a spiny, chalky skeleton and a

water vascular system. Most have a radial body plan
without a head.


Panarthropods with a rigid outer layer and jointed

limbs (arthropod means jointed limb). Arthropods form
by far the most diverse animal group, and examples
include insects, spiders, scorpions, crabs, shrimps,
barnacles, and millipedes.
Species 1,000,000


Species 630






Species 1,500


Ecdysozoans with limbs and claws. This group is

dominated by arthropods, but also includes the small
and obscure velvet worms and tardigrades.


Species 2

Species 1,150



Species 50

Species 940

Species 600

Deuterostomes with a nerve cord along the back and a
rod (notochord) supporting the body. Sea squirts only
have notochords as larvae.

All chordates that possess a skull. Most craniates are
vertebrates. Hagsh are the only invertebrate craniates
although they have a skull, they have no spine.


Species 2,000

Well-developed limbs
characterize amphibians,
reptiles, and mammals.


Membranous sac
An amnion, a waterproof
membrane around a
developing embryo, links
reptiles and mammals
but not amphibians.

Craniates that have a skull and a nerve cord along the

back, and a notochord that develops into a vertebral
column the spine, or backbone. Vertebrates are the
largest animals in most habitats and include sh,
amphibians, reptiles, birds, and mammals.
Species 49,500


Species 50

Color coding
Each major group has a
different color, with closely
related groups in different
shades of the same color.

Where relevant, a crossreference leads on to or
back to related diagrams.

Mammals have a single
bone in the jaw, the
other bones having
become the three bones
of the middle ear.

On the following pages, eight classification diagrams show the
structure of each of the major animal groups. The diagrams, an
example of which is shown above, are based on the evolutionary
relationships used in the phylogenetic system of classification, with
lines and colors tracing the relationships. Each group, enclosed by
a differently colored shape, contains an animal that evolved a new
feature (such as a backbone) plus all of its descendants, most or
all of which inherited that feature. When one of these descendants
evolves a major new feature, a further smaller group containing that
animal and all of its descendants can be defined. This results in a
series of concentric groups, which describes the pattern of both
animal diversity and animal evolution.





Starting with odd-toed ungulates as a whole, tapirs and rhinoceroses
can be separated from horses because of differences in features
of the teeth (character 1). Rhinoceroses have horns on their noses
(character 2) whereas tapirs do not. Only in the two African species
(Diceros and Ceratotherium) does the front horn exceed 3 ft 3 in
(1 m) (character 3). Among the Asian species, the Javan and Indian
rhinoceroses only possess one horn (character 4) whereas the
Sumatran rhinoceros has two, neither of which exceeds 3 ft 3 in
(1 m). The Javan rhinoceros, apart from its different locality, and
appearance, can be isolated on genetic evidence (character 5).



Diceros bicornis
All five species of living rhinoceros have one or two horns.
The Sumatran rhinoceros (top) has two horns, whereas
the Javan rhinoceros (bottom) has one. Genetic evidence
supports treating one horn as the derived condition,
suggesting that the one-horned Javan and Indian species
are more closely related to each other than to the others.






Starting point
Each diagram
starts top left.

Lines and arrows trace the links
between groups. Dotted lines
indicate unresolved relationships.




Animalia Organisms that obtain nutrition from other

organisms and whose bodies are built of many cells,
all of which lack cell walls.

Eumetazoa Animals with integrated multicellular body

structure, with cells organized in tissues. Eumetazoans
include all animals other than sponges, which lack tissues.


Species 20,000

Species 9,000

Species 100

Species 2,000



Species 1

Species 2,000

Species 20,000


Species 150


Species 17,500

Species 17

Bilateria All descendants of eumetazoans with a single

line of symmetry, featuring a front end, usually with
a head. Bilaterians include nearly all animals, even
those that have a radial (wheel-like) arrangement,
such as starsh.

Ecdysozoa Bilaterally symmetrical animals with a threelayered outer covering that is molted. Most ecdysozoans
are arthropods, but roundworms are also abundant.


Species 1,450

Species 90

Species 100

Species 1

Species 1,400

Species 70



Panarthropods with a rigid outer layer and jointed

limbs (arthropod means jointed limb). Arthropods
form by far the most diverse animal group, and
examples include insects, spiders, scorpions, crabs,
shrimps, barnacles, and millipedes.

Panarthropoda Ecdysozoans with limbs or claws.

This group is dominated by arthropods, but also includes
the obscure velvet worms and tardigrades.

Species More than 1,000,000



Species 600


Just over a million animal species have been identied by
scientists, but the total is probably several million. The
vast majority of these are invertebratesanimals without
a backbonefrom simple sponges and jellysh to
sophisticated honey bees. The vast array of invertebrate


Most animals are invertebrates, but this classification
is not labeled on the diagram. This is because they
do not form a cohesive natural group, but many
separate lines of evolution. The area of the diagram
occupied by invertebrates is outlined in blue. Having
evolved from invertebrates, vertebrates also lie within
the blue outline, but have their own discrete section
reflecting their distinctive evolved feature.

groups occupies most of the animal evolutionary tree.

Although relatively few vertebrates have been identied,
they comprise a diverse group ranging in size from tiny
shes less than 3/8 in (1 cm) long to the blue whale, at more
than 98 ft (30 m) long, the largest animal to have ever lived.


Species 8



Species 14,000

Species 35,000



Species 320

Mollusca Lophotrochozoans whose ancestor evolved a

radula (rasping tongue), a muscular foot, and a body
cover called a mantle. Not all mollusks retain these.



Species 1,200

Species 12,000


Lophotrochozoa Bilaterians whose larva (young form) is
of a unique type called a trochophore. This group is
dominated by mollusks but has other important members.



Species 350

Species 650


Solenogastres, Caudofoveates
Species 750


Species 300

Species 20



Species 2,000

Species 4,300

Species 100


Species 1,500

Species 630


Deuterostomia Bilaterians with a radial pattern of cell
division in the early embryo. They include some invertebrates
plus chordates, the group containing vertebrates.


Echinodermata Deuterostomes with a spiny, chalky

skeleton and a water vascular system, but no central nervous
system. Most have a radial body plan without a head.



Species 2

Species 1,150



Species 50

Species 940

Chordata Deuterostomes with a nerve cord along the back
and a rod (notochord) supporting the body. Sea squirts only
have notochords as larvae.

Species 2,000

Craniata All chordates that possess a skull. Most craniates

are vertebrates. Hagshes are the only invertebrate
craniatesthey have a skull, but no vertebrae.

Species 70

Craniates that have a skull and a nerve cord along the

back, and a notochord that develops into a vertebral
columnthe spine, or backbone. Vertebrates are
generally the largest animals in most habitats and
include shes, amphibians, reptiles, birds, and mammals.
Species 49,500



The vast majority of the worlds animals are invertebratesspecies without
a backbone. Vertebratessuch as birds, sh, mammals, reptiles, and
amphibiansmake up less than three percent of all known species.
While vertebrates make up a single group
within the phylum Chordata, invertebrates are
arranged scientically into 30 or so separate
phyla, ranging from simple organisms such as
sponges, atworms, and roundworms to more
complex creatures such as arthropods (see
pp.3841) and mollusks. Asexual reproduction
is widespread among invertebrates, but sexual
reproduction is typical. Hermaphrodites, where
male and female sexual organs occur in the
same individual, are common. In species where

there are separate sexes, the male and the

female do not always have to meet, since
fertilization can take place externally.

of chitin (see p.85). In some groups, body

shape is maintained by means of a tough,
exible cuticle and high internal pressure. One
of the main differences between invertebrate
groups is in body symmetry. Some groups, such
as cnidarians, have radial symmetry, their bodies
being arranged in the same way as the spokes
of a wheel. Invertebrates showing bilateral
symmetry, such as the arthropods, can be
divided down the midline into two equal parts.

Invertebrates do not have bony skeletons, but
many have an internal or external skeleton
of some sort. These skeletons are made from
a variety of materials: hard structures often
consist of crystalline minerals, while the outer
covering (called a cuticle) of arthropods is made


Invertebrate senses can range from simple

systems in sedentary species to complex organs
such as the highly developed eyes of predators.
Many invertebrates can sense dissolved or
airborne chemicals, changes in pressure, gravity,
and portions of the electromagnetic spectrum
including infrared and ultraviolet radiation.

Jellyfish, corals, and

sea anemones

Phylum Porifera

Phylum Cnidaria

Sponges are the simplest of

all living animals and lack true
tissues or organs. They live
attached to solid surfaces,
and feed on nutrients that
they lter from the water using
specialized cells and a system of
canals and pores. The soft parts are
supported by a skeleton of spicules
(slivers of carbon carbonate or silica).

Members of this group of aquatic animals

have radially symmetrical bodies that are
essentially tubes, open at one end. The
tube is either attened into a bell shape
(a medusa) or elongate with the closed end
attached to a hard surface (a polyp). All
cnidarians have tentacles around their
mouths that contain stinging cells for prey
capture and defense. Most species
reproduce asexually by budding. In some
species, such as corals, the new creatures
remain joined to form a colony.

Niphates digitalis 12 in (30 cm)

Bushlike soft coral

Dendronephthya sp.
3ft 3in (1 m) tall

Phylum Platyhelminthes
These worms are simple, bilaterally symmetrical
animals with distinct heads and attened, elongate,
unsegmented bodies. They contain no respiratory
system and no circulatory system. Most species,
such as tapeworms, are parasitic, but some are
free-living predators in freshwater and marine
habitats, and others rely on symbiotic algae.
Atlantic snakelocks anemone
Anemonia viridis Tentacles up to 4 in (10 cm) long

Terrestrial atworm
Bipalium sp.
14 in (35 cm) long

Lions mane jellysh

Cyanea capillata
612 ft (2 m) across

Marine atworm
Pseudobiceros zebra
2 in (5 cm) long

Phylum Nematoda
These small, free-living or parasitic, wormlike
animals are among the most abundant
creatures on the planet. Their unsegmented,
cuticle-covered bodies are round in cross
section and taper toward both ends.
They have no circular muscles
and maintain their shape by high internal
pressure. They move by thrashing their
bodies around in characteristic C- or Sshapes using longitudinal muscle bands.
Intestinal nematode of rats
Nippostrongylus brasiliensis 316 in (4 mm)

Upside-down jellysh
Cassiopea xamachana
12 in (30 cm) across

Tubularian hydroid
Tubularia sp.
6 in (15 cm) tall

Freshwater hydroid
Hydra sp.
1 in (2.5 cm) tall

Starfish, sea urchins,

sea cucumbers

Phylum Annelida

Phylum Echinodermata

The bodies of annelids differ from other worms in

being divided into a series of linked, but partly
independent, functional sections. Each section
contains a set of some of the same organs.
The head is often well developed, with sense
organs, a brain, and a mouth. Locomotion by
expansion and contraction of the body is made
possible by circular and longitudinal muscles.
Parchment worm
Chaetopterus variopedatus
10 in (25 cm)

The bodies of these marine invertebrates are

typically spiny, and usually divided into ve
equal parts arranged symmetrically around a
central point. Their bodies may be drawn out
into arms (as in starsh), feathery (as in sea
lilies), or spherical to cylindrical (as in sea
urchins). A unique internal network, called
the water-vascular system, helps them
move, feed, and exchange gases.

Nereis virens
20 in (50 cm)

Crossaster papposus
10 in (25 cm) across

Bloody Henry
Henricia oculata
6 in (15 cm)

Red cushion star

Porania pulvillus
412 in (12 cm) across

Lumbricus terrestris
10 in (25 cm)

Peacock worm
Sabella pavonina
12 in (30 cm)

Black brittlestar
Ophiocomina nigra
arms up to 1 in (2.5 cm)

Sea cucumber
Pseudocolochirus tricolor
4 in (10 cm)

Medicinal leech
Hirudo verbana
412 in (12 cm)

Northern brittlestar
Ophiothrix fragilis
arms up to 6 in (15 cm)
Edible sea
Echinus esculentus
8 in (20 cm) across

Horse leech
Haemopsis sanguisuga
214 in (6 cm)

Phylum Mollusca
Mollusks have one of the widest ranges of body forms of all
invertebrates, but most species have a head, a soft body mass,
and a muscular foot, which is used for locomotion. In the most
advanced mollusks, the cephalopods, which includes the octopus,
the head is well developed and has sophisticated sense organs,
and parts of the head and foot are modied to form preycapturing arms. All mollusks have one or all of the
following features: a horny, toothed ribbon (the radula)
used for rasping food; a calcium carbonate shell or other
structure covering the upper surface of the body; and a
mantle, which is an outer fold of skin covering the mantle cavity.
Mollusks are adapted to life on land, and in fresh or sea water.

Giant clam
Tridacna gigas
up to 5 ft (1.5 m) across

Queen scallop
Aequipecten opercularis
31 2 in (9 cm)

Cats tongue oyster

Spondylus linguaefelis 3 in (7.5 cm)

Thickened cardita
Megacardita incrassata
11 2 in (4 cm)

Lazarus jewel box

Chama lazarus
3 in (7.5 cm)

Precious turban
Turbo petholatus
214 in (6 cm)

Stepped venus
Chione subimbricata
114 in (3 cm)

Triumphant star turban

Guilfordia triumphans
2 in (5 cm)

Common limpet
Patella vulgata
214 in (6 cm) across

Precious wentletrap
Epitonium scalare
214 in (5.7 cm)


Segmented worms


Snipes bill murex

Haustellum haustellum
5 in (13 cm)

Trapezium horse conch

Pleuroploca trapezium
5 in (13 cm)

Great pond snail

Lymnaea stagnalis
2 in (5 cm)

Imperial harp
Harpa costata
3 in (7.5 cm)

Tent olive
Oliva porphyria
31 2 in (9 cm)

Common snail
Cepaea nemoralis
shell 1 in (2.5 cm) across

Violet snail
Janthina janthina
112 in (4 cm)

The junonia
Scaphella junonia
41 2 in (11 cm)

Tusk shell
Pictodentalium formosum
3 in (7.5 cm)

Polynesian tree snails

Partula sp.
shell 114 in (3 cm)

Giant African snail

Achatina fulicula
12 in (30 cm)

Black slug
Arion ater
6 in (15 cm)

European striped snail

Helix aspersa
shell 112 in (4 cm) across

Marbled chiton
Chiton marmoratus
214 in (6 cm)
Green-blue lettuce sea slug
Elysia crispata
2 in (5 cm)

Sepia ofcinalis
1512 in (40 cm)

Common octopus
Octopus vulgaris
3ft 3in(100 cm)

Blue-ringed octopus
Hapalochlaena lunulata
4 in (10 cm)

Loligo paelei
12 in (30 cm)


This brightly colored neon sea slug
(Nembrotha sp.) is a marine mollusk
of the order Nudibranchia. The slug is
moving among a colony of sea squirts,
on which it feeds.



Page 24

Arthropoda Segmented animals with pairs of jointed limbs

and a rigid exoskeleton. Body segments are aggregated or
fused into functional units, the most universal being a head.

Species 1,000




Species 10,000

Species 2,500

Chelicerata Arthropods with bodies divided into two main

sections. They have no antennae, and the rst pair of limbs
takes the form of pincerlike mouthparts (chelicerae).

Species 4


Species 150

Species 9

Species 13,000

Crustacea Arthropods with three body sections. Their heads
have jaws, two pairs of antennae, and one pair of eyes.
They have two-branched limbs and, typically, gills to breathe.

Myriapoda Arthropods with a head and trunk. The head has

one pair of antennae and simple eyes. Trunk segments have
one pair of legs, but may be fused into double segments.





Species 40,000

Species 1,400

Species 500
Species 32,000


Species 1,000



Species 10,000

Arachnida Mainly terrestrial chelicerates with pairs of

simple eyes. Food is typically liqueed outside the body, and
they breathe through book lungs or tracheae, or both.


Species 2


Species 12,000

Species 5,300

Species 5,000

Malacostraca Includes the larger shrimp- and crab-like
crustaceans. The thorax has an upper shell and eight limbbearing segments; the abdomen has six limb-bearing segments.




Decapoda Eucaridans that use the front three pairs

of limbs for feeding and the back ve pairs for walking.
One pair of walking legs may have pincers.

Eucarida Malacostracans with the upper shell fused to

all segments of the thorax. The eyes of eucaridans are
always stalked.

Species 590


Species 250

Species 25,000

Species 90

Arthropod groups
Arthropods appeared in the seas more than 540 million
years ago. There are four main groups of living species:
chelicerates, myriapods, crustaceans, and hexapods. The
relationship between these groups has been the subject

of debate. For example, the common view that hexapods

and myriapods are most closely related to each other has
been challenged by recent studies, which have suggested
that hexapods are more closely related to crustaceans.


Species 6,000


Species 400

Species 350



Species 5,500



Hexapoda Arthropods with a fused thorax comprising

three segments, each bearing a pair of legs. Their heads
have one pair of antennae.

Insecta A huge group of hexapods with external

mouthparts that are not contained inside a pouch that
can be turned inside out.

Species 370


Species 2,500

Pterygota Insects that have wings in addition to legs.

The second and third segments of the thorax typically
each carry a pair of wings.

Species 800

Neoptera Winged insects in which special muscles and
base joints allow the wings to be folded back along the
body when not in use.




Species 4,000

Species 150

Species 20,000




Neuropterida The least advanced group of

holometabolans with net- or lace-like wing veining
and a simple pupal stage.

Species 2,000

Species 300

Species 25


Species 14

Species 370,000

Species 1,900




Holometabola Neopterans that develop with complete

metamorphosis by means of a pupal stage. Larvae and
adults are dissimilar and have different lifestyles.

Species 2,500

Species 560

Species 3,000




Species 6,000



Species 2,800

Species 2,000

Species 5,000

Species 198,000

Species 82,000

Species 4,000

Species 30

Species 300

Species 165,000

Species 8,000



Species 2,000

Species 122,000

Species 550




strengthened by calcium carbonate, while

in terrestrial species it is topped by a thin layer
of waterproof wax to prevent them from drying
out. As an individual grows, it must shed its
cuticle periodically. Arthropods have a number
of jointed limbs, from three pairs in insects to
many hundreds of pairs in myriapods.

This group of invertebrates forms the largest phylum of all living organisms
and accounts for more than three out of four known species of animal. Their
collective biomass far outweighs that of the vertebrates.
Arthropods share several common features.
All species have a bilaterally symmetrical body
that is divided into segments. Over evolutionary
time these segments have become fused into
functional groups (tagma); one of these tagma
always includes a head. Myriapods have a head
and a body trunk that is made up from a number
of similar segments. In arachnids the head and
thorax are fused, forming a cephalothorax, and the
remaining body segments are fused to form the
abdomen. In insects, a very advanced group of

Internal systems

arthropods, the head is comprised of six fused

segments; a thorax, three segments, and an
abdomen, typically with 11 segments.

All arthropods have an open circulatory system:

their internal organs are bathed in a uid known
as hemolymph, which is moved to the front of
the animal by a dorsal, tubelike heart. Gaseous
exchange is carried out by means of gills,
leaike book lungs, or a branching system of
minute, air-lled tubes called tracheae. The
central nervous system consists of a brain,
situated in the head, connected to paired nerve
cords running along the underside of the body.

The body is covered by the cuticlea tough,
lightweight exoskeleton made of protein and
chitinwhich is secreted by the epidermal cells.
To permit movement, the exoskeleton has joints
and hinges where the cuticle is soft and exible.
In large marine species the exoskeleton is

Sea spiders

Horseshoe crabs

Class Pycnogonida

Class Merostomata

Sea spiders are marine arthropods

with a small, segmented body and
long, slender legs that give them a
supercial resemblance to terrestrial
spiders. All species have a small
head and three trunk segments.

Horseshoe crabs are large,

marine arthropods that are
survivors of a diverse group
that ourished in the seas
300 million years ago.
They have a fused head
and thorax, and six pairs
of limbs, which are covered
by a tough carapace.

Sea spider
Styllopalene longicauda
4 in (7.5 mm)

Horseshoe crab
Limulus polyphemus
up to 2312 in (60 cm)

Class Arachnida
This diverse group of largely terrestrial arthropods
includes spiders, scorpions, harvestmen, ticks, mites,
pseudoscorpions, whip-scorpions, and whip-spiders.
The body is divided into two sections. The head and
thorax are fused to form the cephalothorax, which is
joined to the abdomen, in some species by a narrow
stalk. In scorpions, the abdomen has a tail-like extension
bearing a sting. The cephalothorax carries a pair of
pincerlike chelicerae used for consuming prey, a pair of
limblike pedipalps, and four pairs of normal walking legs.
Arachnids are mostly predatory, although a few are
scavengers and some mites are parasitic. They
have narrow mouths and cannot eat large
pieces of food, so they rely on enzymes
to predigest their prey outside the body
or in a preoral cavity.

Yellow scorpion
Buthus occitanus
11 2 4 in (410 cm)
Chilean scorpion
Centromachetes pococki
1412 in (2.512 cm)

Dactylochelifer sp.
16 316 in (24 mm)

Soft tick
Argas persicus 14 in (6 mm)

Thelyphonus sp.
111 2 in (2.53.5 cm)

Phyrnus sp.
114 11 2 in (34 cm)

House spider
Tegenaria duellica
8 12 in (11.5 cm)

Funnel-web spider
Atrax robustus
4 112 in (24 cm)

Red-kneed tarantula
Brachypelma smithi 4 in (10 cm)

Jumping spider
Salticus sp. 316 516 in (58 mm)


Superclass Crustacea
Crustaceans are a very diverse group, ranging from tiny
copepodsbarely visible to the naked eyeto heavy-bodied
crabs and lobsters. They have two pairs of antennae, compound
eyes on stalks, and a cuticle that is often strengthened with
calcium carbonate. The head and thorax are often covered by
a shieldlike shell or carapace, which extends forward to form
a projection called the rostrum. The thoracic limbs are twobranched and specialized to carry out a range of functions, such
as feeding, locomotion, sensing the environment, and respiration
by mean of basal gills. The rst pair of legs may be enlarged to
form chelipeds with strong claws for food handling, defense, and
signaling. The vast majority of species live in freshwater or the sea.

Brine shrimp
Artemia sp.
8 in (1 cm)

The fire shrimp (Lysmata debelius ) is known to
feed on small parasites and dead skin found
on fish. It can grow up to 2 in (5 cm) in length.

Chthamalus sp.
2 in (1.5 cm)

Antartic krill
Euphausia superba
214 in (6 cm)

Common shrimp
Crangon crangon
312 in (9 cm)

Common prawn
Palaemon serratus
412 in (11 cm)

Spiny spider crab

Maja squinado
carapace 6 in (15 cm)

Brown crab
Cancer pagurus
carapace 12 in (30 cm)

Blue crab
Callinectes sapidus
carapace 7 in (18 cm)

Decorater crab
Naxia tumida
carapace up to 11 2 in (4 cm)

Common hermit crab

Pagurus bernhardus 4 in (10 cm)

Squat lobster
Munida quadrispina
234 in (7 cm)

Common lobster
Homarus gammarus 2312 in (60 cm)

Centipedes and millipedes

Superclass Myriapoda
Myriapods are terrestrial arthropods with biting mandibles and
a single pair of antennae. They are conned to moist habitats
because they do not have a waterproof cuticle. Centipedes are
fast moving and carnivorous with elongate, slightly attened
bodies. The trunk segments each bear a single pair of legs, the
rst pair modied as poison claws used to subdue prey. Millipedes
are mostly herbivorous or scavenging species with elongate,
cylindrical or attened bodies. The rst three segments of the
trunk have no legs, but the remaining segments are fused in pairs,
known as diplosegments, each bearing two pairs of legs. Despite
their common name they never have a thousand legs.

Lithobiid centipede
Lithobius forticatus
4 112 in (0.63.8 cm)

Pill millipede
Glomeris marginata
16 34 in (0.22 cm)

Flat-backed millipede
Polydesmus sp.
16 114 in (0.53.2 cm)


Class Insecta
Insects are the most abundant animals on Earth, and they have
evolved many diverse lifestyles. Although they are mainly
terrestrial, there are a signicant number of aquatic species. The
body of an insect is divided into three main sectionsthe head,
thorax, and abdomen. The head bears external mouthparts, one
pair of antennae, and a pair of compound eyes. The thorax bears
three pairs of legs and, typically, two pairs of wings. The abdomen
contains the major organ systems for digestion and reproduction.
The immature stages of some aquatic species have gills, but all
adult insects breathe air and have a well-developed tracheal
system joined to the outside through
a number of small holes called
spiracles. Insects are the only
invertebrates capable of powered
ight and this, together with their
small size and waterproof cuticle,
has allowed them to colonize a
vast range of habitats.

Jumping bristletail
Dilta littoralis 38 in (1.2 cm)

Lepisma saccharina
16 38 in (0.51 cm)

Siphlonurid mayy
Siphlonurus lacustris 38 in (1 cm)

Southern hawker dragony

Aeshna cyanea
wingspan 4 in (10 cm)

Mountain grasshopper
Stauroderus scalaris
1 in (2.5 cm)

Speckled bush cricket

Leptophyes punctatissima
4 in (2 cm)

Common stoney
Dinocras cephalotes
4 in (2 cm)

Forcula auricularia
8 1 2 in (11.5 cm)

Rottenwood termite
Zootermopsis sp.
8 34 in (12 cm)

American cockroach
Periplaneta americana
111 2 in (2.54 cm)

Praying mantis
Mantis religiosa
4 in (7 cm)

Giant water bug

Lethocerus grandis
314 4 in (810 cm)

Stick insect
Phar nacia sp.
up to 1112 in (29 cm)

Shield bug
Graphosoma italicum
8 in (1 cm)

Thorn bug
Umbonia crassicornis
2 in (1.5 cm)

Bird louse
Menacanthus stramineus
8 316 in (34 mm)

Angamiana aetherea
114 214 in (35.5 cm)

Green leaf insect

Phyllium bioculatum
231 2 in (59 cm)


Xanthostigma xanthostigma
2 in (1.5 cm)

Spoonwing lacewing
Nemoptera sinuata
wingspan 2112 in (55 cm)

Deathwatch beetle
Xestobium rufovillosum
16 3 8 in (29 mm)

Goliath beetle
Goliathus meleagris
314 in (8 cm)

Cyrtotrachelus sp.
2 34 in (1.52 cm)

Leaf beetle
Sagra sp.
1 in (2.5 cm)

Atlas beetle
Chalcosoma atlas
214 in (6 cm)

Leaf weevil
Eupholus schoenherri
1 in (2.5 cm)

House y
Musca domestica
4 516 in (68 mm)

Ctenocephalides felis
16 316 in (25 mm)

Eyed ladybird
Anatis ocellata
8 in (1 cm)

Tachinid y
Formosia moneta
8 in (1 cm)

Horse y
Tabanus sudeticus
1 in (2.5 cm)

Reakirts blue buttery

Hemiargus isola
wingspan 34 114 in (23 cm)

Monarch buttery
Danaus plexippus
wingspan 314 41 2 in (8.512 cm)

Pine emperor moth

Nudaurelia cytherea
wingspan 312 5 in (913 cm)

Bumble bee
Bombus terrestris
2 34 in (1.52 cm)

Giant sphinx moth

Cocytius antaeus
wingspan 57 in (1318 cm)

Jewel wasp
Par nopes car nea
8 12 in (11.5 cm)

German wasp
Vespula germanica
2 in (1 cm)

Basker moth
Euchromia lethe
wingspan 112 2 in
(45 cm)

Wood ant
Formica rufa
16 38 in (0.81 cm)



Page 25

Vertebrata All animals with a vertebral column (backbone

or spine). Vertebrates include all shes except hagshes, plus
mammals, birds, reptiles, and amphibians.



Species 38

Gnathosomata Vertebrates with jaws. They include all land

vertebrates and all shes except lampreys, which are
jawless, and hagshes, which lack jaws and vertebrae.

Bony shes with ray ns built from a fan of narrow bone

or cartilage rods. Because ray-nned shes are so diverse,
they represent the majority of vertebrate species. They
range from tiny guppies to giant sturgeons, and from sea
horses to swordsh.
Species 28,000


Osteichthyes Bony shes and all descendants. They
include all vertebrates with a mineralized skeleton
mammals, birds, reptiles, amphibians, and most shes.


Species 1,080



Chondrichthyes Jawed shes with a skeleton made
of cartilage rather than bone.

Sarcopterygii Vertebrates with either lobed ns

or limbs (tetrapods), which arose from lobed ns.




Species 2

Species 34

Species 6




Under the traditional or Linnaen
classification system, which groups
animals by overall similarity, reptiles
and birds form two separate classes
of vertebrates (see pp.2021). Through
the phylogenetic classification system,
which recognizes the evolutionary
relationships between animals, birds
are grouped within reptiles. This is due
to the fact that they share a common
ancestor with the other groups that
comprise reptiles, including lizards and
snakes, turtles, crocodilans, and tuataras.

Animals with three middle-ear bones and a unique jaw
hinge. Todays mammals also have fur and mammary
glands, but we cannot be sure about the many fossil
forms. The early ancestors of mammals diverged from
reptilelike amniotes more than 200 million years ago.
Species 5,000


Although vertebrates are a very diverse group of animals,
only about 50,000 species have been identieda tiny
fraction of all animal species. The rst vertebrates were
primitive shes, and shes make up more than half of all
living vertebrate species. This evolutionary tree has

tetrapods at its center, and it was the earliest tetrapods that

rst grew limbs and left the water for land. These are the
common ancestors from which the vast array of amphibians,
reptiles, birds, and mammals evolved, populating the land,
taking to the sky, and, in some cases, returning to the sea.


Cold-blooded tetrapods with porous, glandular skin.
Todays amphibians (frogs and toads, salamanders
and newts, and caecilians) are descendants of early
tetrapods that did not develop an amnion (waterproof
egg membrane).
Species 6,000


Species 2

Tetrapoda Bony vertebrates with four limbs or
limblike appendages. The rst land-living vertebrates,
tetrapods include the earliest limbed vertebrate and all
of its descendants, including those that have since lost
their limbs, such as snakes.

Lepidosauria Reptiles that shed skin in large pieces
or as a whole. Lepidosaurs include tuataras, snakes, lizards,
as well as worm lizards (amphisbaenians).

Species 300


Species 7,800

Amniota Tetrapods whose embryo grows inside a
waterproof membrane called an amnion, enabling life
outside bodies of water.

Amniotes with thick skin, horny epidermal (skin surface)

scales, and whose eggs have a mineralized shell. This
group includes warm-blooded birds, and other, scaly, coldblooded reptiles.

Species 17,500


Archosaurs with asymmetrical ight feathers (lost in some
ightless species). Many dinosaurs had feathers, but they
were not ight feathers. More recently evolved birdsnot
including the primitive tinamous and ratiteshave many
more shared features, including a horny, toothless bill,
and a keeled breastbone.

Archosauria Reptiles whose teeth are sunk into sockets.
Their living representatives include only crocodiles and
birds, which are dinosaurs.

Species 23

Species 9,500




Page 36


Actinopterygii Bony shes with ray ns built from a

fan of narrow bone or cartilage rods. This very diverse
group includes more than 28,000 species.

Species 16


Species 27



Species 7

Myxinoidea The only craniates that are not

vertebrates, hagshes have a skull but no vertebral
column (backbone).

Species 1



Petromyzontia The only jawless vertebrates, lampreys

form the sister group to all jawed vertebrates. There are
about 40 species of living lampreys.

Teleostei Ray-nned shes with mobile mouths,

due to moveable upper jaws, and symmetrical
(homocercal) tail ns.

Species 220


Species 856


Species 397


Species 8,000

Euteleostei Modern ray-nned shesthe majority
of teleosts. Each of the other four teleost groups show
various primitive features.


Species 366


Species 400


Species 240

Species 250




Osteichthyes Jawed shes with bony, calcied

skeletons. This group includes all shes except for
hagshes, lampreys, and cartilaginous shes.

Sarcopterygii Bony shes with lobed ns.

These shes were the ancestors of all limbed land
vertebrates (tetrapods).




Species 2

Species 6

Species 38,000

Fish groups
The earliest shes evolved from primitive jawed vertebrates
more than 500 million years ago, and from these, two
main groups emerged. Cartilaginous shes remain largely
unchanged as modern-day sharks, rays and skates, and
chimaeras. Bony shes diverged into lobe-nned shes

and ray-nned shes. The former gave rise to tetrapods,

the rst limbed vertebrates and the ancestors of amphibians,
reptiles, birds, and mammals. Ray-nned shes evolved
into a diverse group that includes more than half of all
vertebrate species, and the vast majority of living shes.


Acanthomorpha Euteleosts with true bony spines in
their dorsal, anal, and pelvic ns.


Species 21

Species 10


Species 1,400

Acanthopterygii Spiny-rayed shes with jaws that can
be protruded when feeding. This group includes about
half of all sh species.

Species 70

Species 315


Species 32

Species 285


Percomorpha The largest and most diverse group
of shes and the most advanced spiny-rayed shes.
Percomorphs share several structural features.


Species 680

Species 360


Species 100

Species 1,500

Chondrichthyes Jawed shes with internal skeletons
made of cartilage, rather than bone.

Species 34

Elasmobranchii The largest group of
cartilaginous shes, including sharks and rays.

Species 400


Species 535

Perciformes Percomorphs with spines and soft rays in
their dorsal and anal ns. They include 10,000 diverse
species of marine and freshwater shes.


Fishes are vertebrate animals that are adapted to live, swim, and breathe
in fresh or salt water. They are the most numerous and diverse vertebrate
animals on Earth and can be found in every conceivable aquatic habitat.
Fishes do not form a natural group. The general
term shes includes four groups of vertebrates
that are as different from one another as
mammals, reptiles, and birds. The largest group
is the bony shes, which includes familiar
species such as cod, salmon, and trout. Sharks,
rays, and chimaeras make up the second group,
the cartilaginous shes. The remaining two small
groups are the jawless marine hagshes and the
lampreys. Land vertebrates (tetrapods) and bony
shes share a common ancestryprimitive bony

shes, the lobe-nned coelacanths, and

lungshes are grouped together with the
tetrapods in modern classication systems.
Lobe-nned shes are the most likely ancestors
of the rst land vertebrates.

Fishes from all these groups typically breathe
using gills to extract oxygen from the water,
swim using ns, are covered with protective
scales or bony plates, and are cold-blooded.

However, there are also lungshes that breathe

air, shes with no scales, and sharks, such as
the salmon shark, that can control their body
temperature. The familiar vertebrate senses of
vision, hearing, touch, taste, and smell are used
by all shes to gather information about their
environment. Most shes have a system of
sensory organs called the lateral line running
along both sides of the body, which detects
vibrations made by other shes and animals
moving through the water. Fins are also
characteristic of most, but not all, shes. These
usually consist of two sets of paired ns (pectoral
and pelvic), one, two, or rarely three dorsal ns,
an anal n, and a caudal (tail) n. The ns are
used for propulsion, maneuverability, and stability.
Some shes, such as the gurnard, even use
their ns to walk along the seabed, and ying
shes use theirs like wings to glide above water.
Lampetra uviatilis
up to 191 2 in (50 cm)

Jawless fishes
Classes Myxini, Petromyzontia
Hagshes and lampreys are long, slimy, eel-like shes with no
biting jaws. Lampreys have a round sucker mouth surrounded
by horny, rasping teeth and are mostly parasitic on other shes.
Hagshes have a slitlike mouth surrounded by four pairs of
tentacles and are mostly scavengers. They do not have a true
backbone; instead, a simple exible rod called a notochord
runs the length of the body.

Myxine glutinosa up to 31 in (80 cm)

Cartilaginous fishes
Class Chondrichthyes
Cartilaginous shes comprise sharks,
rays, and deep-water chimaeras. These
shes have an internal skeleton made
principally of exible cartilage. Males
have a copulatory organ called a clasper
and females give birth to live young or
lay large egg capsules. Special sense
organs, called ampullae of Lorenzini,
allow cartilaginous sh to track other
animals by detecting their electrical elds.
Sharks have strong, replaceable teeth and
rough skin covered in tiny, toothlike dermal
denticles. Skates and rays are at with
winglike pectoral ns and a long, thin tail.
The chimaeras are scaleless with a large
head and ratlike tail.

Leopard shark
Triakis semifasciata
up to 7 ft (2.1 m)

Horn shark
Heterodontus francisci
up to 3 ft 2 in (97 cm)

Blonde ray
Raja brachyura
up to 4 ft (1.2 m)

White-spotted bambooshark
Chiloscyllium plagiosum up to 3 ft 1 in (95 cm)

Electric ray
Torpedo californica
up to 41 2 ft (1.4 m)

Pristiophorus japonicus grows to a length of up
to 5 ft (1.5 m). A pair of barbels on the distinctive
saw-shaped snout are used in combination with
taste sensors to locate prey, such as small fishes
and invertebrates, on the seabed.

Great white shark

Carcharodon carcharias
up to 26 ft (8 m)

Frill shark
Chlamydoselachus anguineus
up to 61 2 ft (2 m)

Spotted ratsh
Hydrolagus colliei
up to 3 ft 1 in (95 cm)

Class Osteichthyes
All bony shes have an internal skeleton of hard, calcied bone,
though in some primitive species this may be part cartilage. With
the exception of the lobe-nned shes, the skeleton extends
into the ns as exible, movable rays and spines. This system
allows the sh to maneuver with far greater precision than sharks
and rays. Most bony shes also have a gas-lled swim bladder,
which is used to adjust their buoyancy and is covered in
overlapping exible scales. While most cartilaginous shes can
be easily recognized as such, bony shes have evolved many
different and sometimes bizarre body shapes and n functions
that enable them to survive in almost every aquatic habitat.

American paddlesh
Polyodon spathula
46 ft (1.21.8 m)

Latimeria chalumnae is 56 ft (1.51.8 m)
long. It is found on steep, rocky reefs at depths
of 5002,300 ft (150700 m).

Longnose gar
Lepisosteus osseus
up to 514 ft (1.5 m)
Freshwater butterysh
Pantodon buchholzi
up to 4 in (10 cm)

Barred bichir
Polypterus delhezi
up to 171 2 in (44 cm)

Clown knifesh
Notopterus chitala
up to 4 ft (1.2 m)

Longnosed elephant sh
Gnathonemus petersii
up to 14 in (35 cm)

Megalops atlanticus
up to 41 2 814 ft (1.32.5 m)

Jewel moray eel

Muraena lentiginosa up to 231 2 in (60 cm)

Ringed snake eel

Myrichthys colubrinus
up to 35 in (88 cm)

Sea gulper eel

Eurypharynx pelecanoides
2312 39 in (60100 cm)

Blue ribbon eel

Rhinomuraena quaesita up to 414 ft (1.3 m)

Elegant corydoras
Corydoras elegans
up to 214 in (6 cm)

Chanos chanos
up to 6 ft (1.8 m)
Black angel catsh
Synodontis angelicus
up to 2112 in (55 cm)

Walking catsh
Clarias batrachus
up to 211 2 in (55 cm)

Red-bellied piranha
Pygocentrus nattereri
up to 12 in (30 cm)

Silver hatchetsh
Gasteropelecus sternicla
up to 214 in (6 cm)


Bony fishes


Rust-colored rudd
Scardinius erythrophthalmus
up to 4 in (10 cm)

Rainbow trout
Oncorhynchus mykiss up to 4 ft (1.2 m)

Marine hatchetsh
Argyropelecus aculeatus
up to 212 in (7 cm)

Northern pike
Esox lucius up to 414 ft (1.3 m)

Atlantic cod
Gadus morhua
up to 612 ft (2 m)

Sockeye salmon
Oncorhynchus nerka
up to 33 in (84 cm)

Berndts beardsh
Polymixia berndti
up to 1814 in (47.5 cm)

Splendid toadsh
Sanopus splendidus up to 912 in (24 cm)

Sargassum sh
Histrio histrio
up to 7 in (18.5 cm)

Crimson soldiersh
Myripristis murdjan
up to 231 2 in (60 cm)

Threadn rainbowsh
Iriatherina werneri
up to 112 in (4 cm)

Silver needlesh
Xenentodon cancila up to 12 in (30 cm)

John Dory
Zeus faber
up to 35 in (90 cm)

Madagascar rainbowsh
Bedotia geayi
up to 4 in (10 cm)

Three-spined stickleback
Gasterosteus aculeatus
up to 31 2 in (8 cm)

Sea horse
up to 6 in (15 cm)

Weedy seadragon
Phyllopteryx taeniolatus
up to 18 in (46 cm)

Talibar lionsh
Pterois radiata up to 10 in (25 cm)


Reef stonesh
Synanceia verrucosa
up to 1512 in (40 cm)

Tub gurnard
Chelidonichthyes lucernus
up to 30 in (75 cm)

Harlequin tusksh
Choerodon fasciatus
up to 10 in (25 cm)

Black chipokae cichlid

Melanochromis chipokae up to 5 in (13 cm)

Queen angelsh
Holacanthus ciliaris
up to 171 2 in (45 cm)

Black-nned butterysh
Chaetodon decussatus
up to 8 in (20 cm)

Rock cock
Centrolabrus exoletus
up to 61 2 in (17.5 cm)

European perch
Perca uviatilis
up to 20 in (51 cm)

Threadn butterysh
Chaetodon auriga
up to 8 in (20 cm)

Maroon clownsh
Premnas biaculeatus up to 6 in (15 cm)

Polka-dot grouper
Chromileptes altivelis up to 24 in (60 cm)

European ounder
Pleuronectes platessa
up to 3 ft 3 in (1 m)

Atlantic halibut
Hippoglossus hippoglossus
up to 1434 ft (4.5 m)

Silver sailn molly

Poecilia latipinna up to 4 in (10 cm)

Mandarin sh
Synchiropus splendidus
up to 3 in (7.5 cm)

Spotted trunksh
Lactophrys triqueter
up to 1812 in (47 cm)

Amiets killish
Fundulopanchax amieti up to 234 in (7 cm)

The ocean sunfish (Mola mola ) is up
to 13 ft (4 m) in length, and is the heaviest
bony fish, weighing up to 2.2 tons.

Dwarf Argentine pearlsh

Austrolebias nigripinnis up to 2 in (5 cm)

Tasselled lesh
Chaetodermis pencilligerus
up to 10 in (25 cm)




Amphibia Cold-blooded tetrapods. Most produce eggs

that hatch into larvae then metamorphose into adults. Their
porous and glandular skin is kept moist.

Batrachia All amphibians other than the limbless

caecilians. The group includes all types of newts and
salamanders and frogs and toads.

Page 37





Anura Tailless amphibians usually with large hind limbs

adapted for jumping and swimming. Most lay eggs that
hatch into tadpoles, which metamorphose into adults.

Lalagobatrachia All frogs other than tailed and

New Zealand frogs. They communicate by calling
and most have vocal sacs.

Species 1




Species 4

Species 31

Gymnophiona Limbless, wormlike amphibians that

burrow in soil or live in rivers. Some species lay eggs,
producing larvae; others give birth to live young.

Species 11

Species 10

Species 164




Species 505

Species 146


There is no clear distinction between frogs
and toads. Together they make up a group
called the anurans. True toads are members
of a smaller group, called the Bufonidae, but
the name toad is often used for any anuran
that has dry, warty skin and spends most of
its life on land. Frogs have smooth, moist skin
and spend a lot of their time in water. Most
anurans undergo a radical metamorphosis
from aquatic tadpoles to terrestrial adults.



Species 844

Species 97

Amphibian groups
Amphibians have existed for at least 230 million years,
when they evolved from shes. It is not clear whether the
three main groupscaecilians, salamanders and newts,
and frogs and toadsare descended from a common
ancestor or whether they evolved from different groups.

Amphibians are often erroneously seen as an intermediate

group between shes and reptiles. In fact they have evolved
adaptations for living in moist habitats around freshwater.
This diagram represents a recent hypothesis of amphibian
relationships, and not all families are shown here.




Caudata Amphibians with elongated bodies, short legs,

and long tails. Fertilization is mostly internal and most lay
eggs, but some give birth to larvae or small adults.

Species 51


Cryptobranchoidea Salamanders with external
fertilization. This group includes giant salamanders and
a group of primitive Asiatic salamanders.

Species 4



Species 3

Salamanders with internal fertilization carried out using

a spermatophore (sperm capsule) that is transferred
from the male into the female.


Species 378

Species 3

Species 37

Treptobranchia Salamanders that as larvae are aquatic
with external gills, and as adults are terrestrial and
breathe using lungs.


Species 74

Species 6

Species 6

Perennibranchia Permanently aquatic salamanders
with bushy external gills both as larvae and adults.


Species 4

Species 426



Species 286

Species 207

Species 166

Species 315


Species 132


Species 64


Amphibians are ectothermic (cold-blooded) vertebrates that derive body heat
from their environment. They have four limbs, smooth skin, and are divided
into three groups: caecilians, newts and salamanders, and frogs and toads.
Amphibians are typically aquatic as larvae and
terrestrial as adults. They are highly dependent
on water and are associated with freshwater
habitatsnone live in the sea. They typically
produce eggs that lack shells to prevent water
loss and have to be laid in water. Eggs hatch into
larvae (called tadpoles in frogs and toads) that
live in water for an extended period. Larvae
undergo a complex transformation, called
metamorphosis, in which a number of changes
in body form equip them for life as adults in

terrestrial habitats. Most importantly, the larval

gills are lost and replaced by air-breathing lungs.
In frogs and toads, the tail is reabsorbed during
metamorphosis and the limbs develop.

The skin of amphibians contains glands, is
commonly moist, and lacks scales, feathers,
or hair. Most amphibians have lungs but all, to a
greater or lesser extent, take in oxygen through
their skin. The skin is also important for taking in

water from the environment and contains

numerous pigment cells that give it color. Many
amphibians are brightly colored; this is especially
common in poisonous species, in which striking
colors and skin patterns serve as a warning.
In other species, skin coloration provides
camouage or plays a role in mate selection.

Parental care
Amphibians have diverse life histories. While
many produce very large numbers of eggs that
they do not care for, others have evolved parental
care in which a small number of eggs are
protected by one or both parents. Parental care
is usually provided by the female in caecilians,
newts, and salamanders, but in many frogs it is
provided by the male. Lacking a shell, amphibian
eggs need protection against dehydration and
infection by fungae, as well as from predators.

Order Gymnophiona
With a total of 175 species in three families, caecilians form
the smallest of the three major groups of amphibians. They
are limbless, wormlike animals that burrow in the soil in
forested habitats, although some species live in rivers and
streams. Conned to tropical habitats in South America,
Africa, and Southeast Asia, caecilians vary in length from
several centimetres to more than 1.5m (5ft). Fertilization is
internal. Some species lay eggs, from which free-living larvae
emerge, while in others development through metamorphosis
occurs within the mother, who gives birth to small adults.
Caecilians are difcult to observe and their biology is less
well known than that of other amphibians.


Ichthyophis glutinosus, seen protecting its eggs, is
1215 1 2 in (3040 cm) long. It lives underground
and feeds on worms and other invertebrates.

Ringed caecilian
Siphonops annulatus
8151 2 in (2040 cm)

Salamanders and newts

Order Caudata
These tailed amphibians generally lead
secretive lives. They live in cool, shady
places and are typically active at night.
Absent from most of Africa and Australia,
salamanders and newts occur in temperate
and tropical habitats in North and South
America, Europe, and Asia. There are 563
species in nine families, and few are more
than 6 in (15 cm) long. Fertilization is usually
internal. Most species lay eggs but in some
the eggs hatch inside the mother, who gives
birth to larvae or small adults. The male
lacks a penis and sperm is transferred
to the female in a special structure called
a spermatophore. Some species
are aquatic, others are
wholly terrestrial, and
others spend their
lives partly in water
and partly on land.

Necturus maculosus 8191 2 in (2050 cm)

Cryptobranchus alleganiensis
1230 in (3075 cm)

Fire salamander
Salamandra salamandra
314 11 in (828 cm)

Mexican axolotl
Ambystoma mexicanum
48 in (1020 cm)

Californian newt
Taricha torosa
41 2 8 in (1220 cm)

Himalayan newt
Tylototriton verrucosus
41 2 7 in (1218 cm)

One of the longest amphibians in North America,
Siren lacertina is 19 1 2 35 in (5090 cm) long.
This species has only one pair of small legs,
situated behind the feathery external gills.

Great crested newt

Triturus cristatus
46 in (1015 cm)

Pacic giant salamander

Dicamptodon tenebrosus
612 1312 in (1734 cm)


Frogs and toads

Order Anura
Frogs and toads lack tails, at least as adults. Their hind limbs
are much larger than their front legs and are adapted for jumping,
swimming, and digging. Most are active at night and communicate
by calling. Frogs and toads occur in temperate and tropical
habitats in all continents. There are 5,572 species in 44 families
and few are more than 14 in (35 cm) long. Fertilization is external.
Most species lay eggs that hatch into free-swimming tadpoles,
which then metamorphose into small adults. In some the eggs
develop in or on one of the parents. Some species spend their
lives partly in water and partly on land, some are aquatic
throughout their lives, while others are wholly terrestrial.
Tailed frog
Ascaphus truei
12 in (2.55 cm)

Mexican burrowing toad

Rhinophrynus dorsalis
21 2 314 in (68 cm)

African clawed frog

Xenopus laevis
214 5 in (613 cm)

Midwife toad
Alytes obstetricans
114 2 in (35 cm)

Mallorcan midwife toad

Alytes muletensis
114 134 in (34.5 cm)

Oriental re-bellied toad

Bombina orientalis
114 2 in (35 cm)

Parsley frog
Pelodytes punctatus
114 2 in (35 cm)

Couchs spadefoot toad

Scaphiopus couchii
214 31 2 in (5.59 cm)

Asian leaf frog

Megophrys montana
234 51 2 in (714 cm)

Cape ghost frog

Heleophryne purcelli
114 214 in (36 cm)

Seychelles frog
Sechellophryne gardineri
8 1 2 in (11.5 cm)

Sign-bearing froglet
Crinia insignifera
2 114 in (1.53 cm)

Mountain marsupial frog

Gastrotheca monticola
112 214 in (46 cm)

Whites treefrog
Litoria caerulea
24 in (510 cm)

Red-eyed treefrog
Agalychnis callidryas
112 234 in (47 cm)

Green treefrog
Hyla cinerea
114 214 in (36 cm)

European treefrog
Hyla arborea 114 2 in (35 cm)

Surinam horned frog

Ceratophrys cornuta 48 in (1020 cm)


Green poison frog

Dendrobates auratus
1214 in (2.56 cm)

Green toad
Pseudepidalea viridis
31 2 412 in (912 cm)


The electric-blue coloration of Dendrobates
tinctorius warns predators of its highly poisonous
nature. It is 11/ 4 2in (35 cm) long and is found in
the tropical forests of South America.

Marine toad
Rhinella marina
4912 in (1024 cm)

European common toad

Bufo bufo 314 8 in (820 cm)

Tomato frog
Dyscophus antongilii
314 412 in (812 cm)

Panamanian golden frog

Atelopus zeteki
112 214 in (45.5 cm)

Yellow-banded poison frog

Dendrobates leucomelas
114 11 2 in (33.5 cm)

Boulengers Asian tree toad

Pedostibes hosii
24 in (510 cm)

African bullfrog
Pyxicephalus adspersus
334 9 in (823 cm)

Kassina frog
Kassina senegalensis 114 2 in (35 cm)
Tinker reed frog
Hyperolius tuberilinguis
114 134 in (34.5 cm)

Green mantella
Mantella viridis 34 114 in (23 cm)

Golden mantella
Mantella aurantiaca 34 114 in (23 cm)

Solomon Island horned frog
Ceratobatrachus guentheri
2314 in (58 cm)

European common frog

Rana temporaria
24 in (510 cm)

Agile frog
Rana dalmatina
231 2 in (59 cm)

Lithobates catesbeianus is the largest frog in North

America, growing to a length of up to 8 in (20 cm). It
lives in lakes, ponds, and slow-flowing streams.



Found in the rain forests of Costa Rica,
Nicaragua, and Panama, the strawberry
poison frog (Oophaga pumilio) is
4 1 in (22.5 cm) long and may
exhibit remarkable color variation.




Reptilia Thick-skinned tetrapods with amniotic eggs or

internal development of young. They include warm-blooded
birds and other scaly, cold-blooded reptiles.

Diapsida All reptiles, other than turtles, including birds.

Body forms vary but are often elongated, with scaly or
feathered coverings and, commonly, four limbs.

Page 37

Species 2

Lepidosauria Diapsids that shed their skin in large pieces
or as a whole.

Species 255

Testudines Reptiles with bodies contained within
upper and lower bony, boxlike shells. There are aquatic,
semiaquatic, and terrestrial species.



Species 38

Species 398


Iguania Lizards that have four functional limbs and use
their tongues to capture and grab food.

Species 54



Archosauria Reptiles with teeth sunk into sockets. This

group includes many extinct dinosaur groups, with birds and
crocodilians being the only living archosaurs.


Species 567

Species 570

Archosaurs with asymmetrical ight feathers (lost in
some ightless species). Many dinosaurs had feathers,
but they were not ight feathers. More recently evolved
birdsnot including the primitive tinamous and
ratiteshave many more shared features, including a
horny, toothless bill and a keeled breastbone.
Species 9,500

Crocodylia Elongated, limbed reptiles covered with thick
leathery plates under which are bony plates on their top
surface. All are semiaquatic predators.



Lizards and snakes are closely related
and make up the group Squamata. It is
thought that snakes evolved from lizards,
possibly from burrowing species, and the
differences between them are slight.


Reptile groups
The oldest groups of reptiles are the turtles and archosaurs,
which rst appeared about 220 million years ago.
Archosaurs include crocodilians and birds, which are
feathered reptiles, but also the extinct dinosaurs.
Lepidosaurs include the tuataras, relics of a once




widespread group of lizardlike reptiles, and the squamates, a

huge group that includes all lizards and snakes. This
diagram represents one hypothesis of reptile relationships.
Controversy surrounds the relations between squamates
and the placement of turtles. Not all families are shown here.




Squamata Scaly reptiles of many forms, also known

as squamates. Males have paired copulatory organs
called hemipenes.


Species 166

Species 1,073

Scleroglossa Squamates that use their jaws rather than
their tongue to catch food, and their tongue for smelling.
This group includes all squamates except iguanians.


Species 2,000

Autarchoglossa Squamates with highly developed
olfactory capabilities involving the tongue and a sensitive
scent organ (Jacobsons organ) in the roof of the mouth.



Species 6

Species 114

Anguimorpha A diverse group of squamates. Many have
bony plates (osteoderms) beneath their scales
and some species lack legs.



Species 2

Species 63


Varanoidea Includes squamates with an excellent sense of
smell, and well-developed teeth, upper jaw, and neck. Most
are efcient predators that use long tongues to track prey.

Serpentes Elongated squamates with no limbs (although
the skeletons of some snakes show evidence of hind
limbs), no moveable eyelids, and no external ears.

Species 473

Species 2,500


higher temperatures than endothermic (warmblooded) birds and mammals. Some reptiles
may also survive temperatures as low as
freezing, although in these conditions their
bodily functions operate at a very reduced rate.

What most people understand by the term reptile is an egg-bearing
vertebrates that has a tough skin with a covering of hard, dry scales.
In fact, the term also includes birds, with their covering of feathers.
Living reptiles include turtles and tortoises,
tuatara, squamates (lizards, amphisbaenians,
and snakes), crocodilians (crocodiles, alligators,
and caimans), and, perhaps somewhat
surprisingly, birds. In fact, crocodilians are more
closely related to the birds than to the other
reptiles, and together they form a group known
as the Archosaura (ruling reptiles), which also
contains several extinct groups of dinosaurs and
pterosaurs (ying reptiles). Birds, then, are
warm-blooded reptiles with feathers. For the

Extreme habitats

purposes of this catalog, the birds have been

proled separately from the non-avian reptiles
(see pp.5867)

Temperature control
Reptiles occur throughout the world but are
more common in tropical habitats. Most nonavian reptiles are ectothermic, meaning they
cannot generate metabolic heat. However, some
regulate their internal body temperature through
behavior, such as sunning, and can operate at

By obtaining all their heat from external sources

such as the sun, ectothermic reptiles can
survive easily on one-tenth of the amount
of food needed by an endothermic animal
of the same sizeendotherms use up a large
proportion of the energy obtained from their
food just to maintain a constant body
temperature. This gives non-avian reptiles a
great advantage in habitats where conditions
are extreme and food is in short supply, notably
deserts, where lizards are often the most
common form of vertebrate life.

Turtles and tortoises

Order Chelonia
Turtles and tortoises comprise the order Chelonia.
They are heavily armored reptiles with bony shells that
originate from their ribs and cover the top and bottom
of their bodies. This gives them excellent protection
from predators but limits their speed and agility on land.
They all lack teeth and instead use a sharp horny beak
to break up their food, which may consist of animal or
vegetable material, or both, depending on the species. Turtles
occur on the land (where they are often known as tortoises), in
freshwater, and in the sea. They all lay nearly spherical eggs,
with some marine species laying up to 100 in a clutch.

Alligator snapping turtle

Macroclemys temminckii
1512 31 in (4080 cm)

Big-headed turtle
Platysternon megacephalum
512 8 in (1420 cm)

Green sea turtle

Chelonia mydas
31 in3 ft 3 in (80100 cm)

Starred tortoise
Geochelone elegans
1215 in (3038 cm)

Red-eared turtle
Trachemys scripta elegans
811 in (2028 cm)

Hermanns tortoise
Testudo hermanni
68 in (1520 cm)

Yellow mud turtle

Kinosternon avescens
41 2 612 in (1216 cm)

Leopard tortoise
Geochelone pardalis
17 12 28 in (4572 cm)

Chinese soft-shelled turtle

Pelodiscus sinensis
612 in (1530 cm)

Order Rhynchocephalia
Both surviving members of the order Rhynchocephalia live
on islands off the coast of New Zealand. Despite their lizardlike
appearance, they are not squamates and belong to an ancient
group of reptiles that was formerly spread over much of the world.
Their teeth are fused to their jawbones and those on the tip of their
upper jaw are modied into a beaklike arrangement. Tuataras also
lack the hemi penis (double penis) unique to squamates. They lay
eggs, which take about a year to hatch, and have long life spans.

Sphenodon punctatus
1912 26 in (5065 cm)

Common snake-necked turtle

Chelodina longicollis
810 in (2025 cm)


Lizards and snakes

Order Squamata
The lizards, snakes, and amphisbaenians (worm-lizards)
comprise the Squamata. This is the largest group of ectothermic
reptiles, with the widest distribution. Most squamates lay eggs
but a sizable proportion bear live young. Lizards are the most
numerous group and are distinguished by the presence
of legs, although limblessness has evolved many times
among squamates. They also have eyelids and external
ear openings. Squamates have adapted to many
habitats and lled many ecological niches. The wormlizards are burrowing squamates restricted to warm
climates. Their scales are arranged in annuli (rings) and
their skulls are bony, allowing them to force their way
through hardened soil. Most worm-lizards have no limbs,
but the ajolotes or mole lizards (Bipedidae) have front limbs
that they use to dig. Snakes are elongated squamates
lacking legs, eyelids, and external ear openings. These
constraints have resulted in specialized methods of
locomotion and hunting. All snakes are carnivorous,
feeding on animals ranging from ants to antelopes.

Spiny dab lizard

Uromastyx acanthinurus
12151 2 in (3040 cm)

Thorny devil
Moloch horridus
67 in (1518 cm)

Australian frilled lizard

Chlamydosaurus kingii
2312 35 in (6090 cm)

Inland bearded dragon

Pogona vitticeps
121912 in (3050 cm)

Warty chameleon
Furcifer verrucosus
410 in (1026 cm)

Common agama
Agama agama
121512 in (3040 cm)

Thai water dragon

Physignathus cocincinus
3139 in (80100 cm)

Flying lizard
Draco spilonotus
68 in (1520 cm)

Jacksons chameleon
Chamaeleo jacksonii
812 in (2030 cm)
Plumed basilisk
Basiliscus plumifrons
231 2 28 in (6070 cm)

Collared lizard
Crotaphtus collaris
814 in (2035 cm)

Desert horned lizard

Phrynosoma platyrhinos
314 41 2 in (811 cm)

Marine iguana
Amblyrhynchus cristatus
3ft 3in512 ft (11.7 m)

Green iguana
Iguana iguana
5612 ft (1.52 m)

Green anolis
Anolis carolinensis
412 8 in (1220 cm)


Tokay gecko
Gekko gecko
714 in (1835 cm)

Kuhls ying gecko

Ptychozoon kuhli
78 in (1820 cm)

Rough-scaled plated lizard

Gerrhosaurus major 1619 in (4148 cm)

Northern leaf-tailed gecko

Phyllurus cornutus
681 2 in (1521 cm)

Nicobar blind lizard

Dibamus nicobaricus 45 in (1013 cm)

Tegu lizard
Tupinambis teguixin
3139 in (80100 cm)

Yellow-spotted night lizard

Lepidophyma avimaculatum
812 in (2030 cm)

Eyed skink
Chalcides ocellatus
712 in (1830 cm)

Armadillo lizard
Cordylus cataphractus
61 2 81 2 in (1621 cm)

Ocellated lizard
Lacerta lepida
2312 31 in (6080 cm)

European glass lizard

Ophisaurus apodus
3ft 3in4 ft (11.2 m)

Savanna monitor
21 2 4 ft (0.81.2 m)

Black-and-pink-striped gila monster

Heloderma suspectum
12191 2 in (3050 cm)

Komodo dragon
Varanus komodoensis
814 1014 ft (2.53.1 m)

Amphisbaena fuliginosa
1217 12 in (3045 cm)

Leptotyphlops dulcis grows to a length of 610 12 in (1527 cm).
It is a member of the Leptotyphlopidae, a family of small,
burrowing snakes that live in the soil and feed mainly on termites.


Anilius scytale is 2835 in (7090 cm) long. The sole
member of its family, its markings mimic those of
venomous coral snakes that live in the same region.


Sunbeam snake
Xenopeltis unicolor
3ft 3in4 ft (11.3 m)

Eunectes murinus
2033 ft (610 m)

Arafura lesnake
Acrochordus arafurae
5814 ft (1.52.5 m)

Rosy boa
Charina trivirgata
2312 ft (0.61.1 m)

Mexican burrowing snake

Loxocemus bicolor
3ft 3in4 ft (11.3 m)

Burmese python
Python molurus
1612 23 ft (57 m)

Gaboon viper
Bitis gabonica 4612 ft (1.22 m)

Emerald tree boa

Corallus caninus
561 2 ft (1.52 m)


Crotalus atrox is a large and formidable pit viper,
37 ft (12.1 m) long, common in the arid American
Southwest. It is active mainly at night.

Puff adder
Bitis arietans
36 ft (0.91.8 m)

Egyptian cobra
Naja haje
5734 ft (1.52.4 m)

Elaphe guttata
36 ft (11.8 m)

Bibrons burrowing asp

Atractaspis bibroni
2028 in (5070 cm)

Cuban woodsnake
Tropidophis melanurus 31 in39in (80100 cm)

Eastern coral snake

Micrurus fulvius
2839 in (70100 cm)

Crocodiles, alligators, and gharials

Order Crocodilia
Crocodilians are one of the two surviving groups from an
evolutionary line that also contained the extinct dinosaurs, the
other survivors being the birds. Crocodilians are covered with
thick bony plates and are semiaquatic predators. Their social
behavior, complex displays, and vocalizations set them apart
from all other reptiles. All species lay eggs, and females show
a high degree of parental care. Most crocodilians live in
freshwater rivers, lakes, and lagoons, while a few species
inhabit tidal reaches and may venture out to sea.

American alligator
Alligator mississippiensis
914 161 2 ft (2.85 m)

Nile crocodile
Crocodylus niloticus
161 2 21 ft (56.5 m)

Spectacled caiman
Caiman crocodilus
814 934 ft (2.53 m)

Gavialis gangeticus
1323 ft (47 m)




Aves Feathered vertebrates with forelimbs modied into

wings (in most cases), and a horny toothless bill. Birds also
lay hard-shelled eggs.

Neognathae All living birds except the tinamous and the

ightless ratites. A huge variety of forms has evolved,
including the passerines (perching birds).

Page 37

Species 298

Species 16



Species 46

Species 429




Galliformes Largely terrestrial birds, with short, stout,

decurved bills, and strong legs and feet. Short wings
typically give short bursts of low ight. This group includes
the jungle fowl, from which domestic fowl were derived.

Species 4

Species 1

Piciformes Neoaves with feet having two toes pointing

forward and two backward. They have chisel-like bills, and
many have stiff tail feathers acting as a prop when perched
on upright branches.

Ratitae Flightless birds with reduced ight muscles, no

developed ight feathers, and a breastbone with no keel.
This group includes the largest living birds.


Species 2

Species 115



Species 6



Anseriformes Water or waterside birds with three toes
on each foot joined by webs. Their bills vary from slender
and serrated to broad and triangular. This group includes
ducks, geese, and swans.

Species 6

Species 39

Coraciiformes Tiny to medium-sized, dry-land
or waterside birds, with two toes fused at the base,
typically a short, square tail, an upright stance, and
a horizontal bill. Many have striking colors.

Bird groups
Birds number close to 10,000 species, ranging from the
primitive tinamous and ratites to perching birds, the most
recently evolved group. Outward appearances make some
groupings such as penguins and hummingbirds obvious,
but birds within other groups, such as the cranes and

rails, may appear to have little in common. Recent DNA

studies have helped resolve some of the relationships,
producing dramatically revised groupings. Surprises have
included the close relationship between New World
vultures and storks, rather than other birds of prey.






Species 22

Neoaves All neognaths except the fowl and the

waterfowl. The relationships between groups within
the neoaves remain largely unresolved.

Species 5


Species 60


Pelicaniformes Relatively large sea- and freshwater
birds, with long, angular wings, a bare and exible throat
pouch, a dagger-shaped or hooked bill, forward-facing
eyes, and webbing joining all four toes.


Species 225

Species 199

Species 352



Species 138

Charadriiformes Shore and seabirds, with bill shape

and leg length adapted to their feeding techniques. Feet
may be fully or partially webbed, or unwebbed. Found
worldwide, they include long-distance migrants.

Species 23


Species 5


Species 17

Species 107

Species 1

Species 194



Passeriformes Birds with four unwebbed toes joined at

the heel, three pointing forward and one backward. Also
known as passerines, they vary hugely in form.

Tyranni A group of South American passerines, divided

from songbirds by physical features including the detailed
structure of the vocal organ (syrinx).



Species 12

Species 3



Species 174

Species 2

Species 2

Corvidae Passerines with strong, heavily scaled feet and
a stout, versatile bill. With loud and hoarse calls, they are
often social birds and highly developed mentally. This
group includes crows, jays, and magpies.

Passeri A group of passerines, ranging greatly in size, with

a complex vocal organ (syrinx) that gives many species a
sophisticated vocal range and control.

Species 5

Species 7

Species 3

Species 18

Passerida A group of songbirds comprising an

extremely varied mixture of species. Some subgroups are
very distinct but the true relationships between the
different groups are yet to be resolved. They include
about one-third of the worlds bird species.



the tracts are harder to dene. Feathers provide

insulation, enable ight, and need to be replaced
regularly through the process of molting,
usually once or twice a year.

Birds are lightweight but remarkably tough animals, with a high metabolic rate
and often fast, high-energy lifestyles. They are found throughout the world,
except for the most extreme polar areas, and in nearly every surface habitat.

Counting toes

Modern birds trace their descent back to dinosaurs

and have much in common with reptiles, but all
bird species are warm-blooded. Most species
can y, but there are some ightless ones. All
species lay eggs that are incubated externally,
and have horn-sheathed bills. There is, however,
another feature that is unique to birds: the

feather. Feathers developed from modied scales,

and grow in well-dened tracts. On most birds
these tracts form a regular pattern, described by
such terms as ear coverts, scapulars, primaries,
and tail feathers. Overlying the body, smaller
contour feathers smooth the birds outline. On
some species, including penguins and kiwis,

By using detailed physical and behavioral

features to trace common descent, birds
are grouped into 227 families to create an
evolutionary family tree. Physical characters
include the structure of the feet: most birds
have four toes, the majority having three facing
forward, one back. A substantial number of
species have just three toes and a few,
including the ostrich, have just two toes.




Cassowaries and emus

Order Tinamiformes

Order Struthioniformes

Order Rheiformes

Order Casuariiformes

This ancient South American family of

ground-living, quail-like birds lives mostly
in dense, tropical forest as far north as
Mexico. Some species live in open
grasslands. Elusive but often
heard, they eat insects, seeds,
and berries, and range from
6191 2 in (1550 cm)
in length.

The worlds largest birds are

ightless and can run at
up to 40 mph
(65 kph).

Rheas are ightless, polygamous

birds that are ostrichlike in form
but have three toes and are
considerably smaller, up to 5 ft
(1.5 m) tall. They live in open
habitats in South
America. They
eat leaves,
shoots, seeds,
and some

These ightless birds, found in

Australia and New Guinea, are
somewhat ostrichlike, but with
a longer, lower prole and
three toes. Cassowaries can
reach 6 ft (1.8 m) in height
and emus, the worlds
birds, can reach
61 2 ft (2 m) and
weigh up to
100 lb (45 kg).

Struthio camelus
7912 ft (2.12.8 m)

Greater rhea
Rhea americana
35 ft (0.91.5 m)

Southern cassowary
Casuarius casuarius
6 ft (1.8 m)

Red-winged tinamou
1512 in (40 cm)


Gamebirds and relatives

Order Apterygiformes

Order Galliformes

Three species of small ightless birds make up

this order. Females are larger than males, and
their especially large eggs are each up to 25
percent of the females body weight. They are
nocturnal, and nd food, especially earthworms,
by touch and scent, using facial bristles and nostrils
at the tip of a long bill.

Gamebirds occur in many habitats,

including semi-desert, grassland,
savanna, woodland and forest, and
even high peaks and northern tundra.
They are all small-headed, large-bodied
birds with short, stout, arched bills and
often marked sexual differences. Some
cold-climate species have feathered
feet and turn white in winter.

Brown kiwi
Apteryx australis

1514 in (40 cm)

California quail
Callipepla californica
10 in (25 cm)

Alectoris chukar
121 2 151 2 in (3239 cm)

Order Anseriformes
This group consists of ducks, geese,
swans, and screamers. Wildfowl are
mostly water or waterside birds,
many of which feed on dry land
and retreat to water for safety.
Swans are the largest, geese mostly
intermediate, and ducks smaller. Ducks
can be freshwater, marine, or both, and
feed on land and in water.
Eurasian wigeon
Anas penelope
3034 in (7586 cm)

Plumed whistling-duck
Dendrocygna eytoni 151 2 1712 in (4045 cm)

Northern bobwhite
Colinus virginianus
912 11 in (2428 cm)

Western capercaillie
Tetrao urogallus
231 2 33 in (6085 cm)

Mute swan
Cygnus olor
412 514 ft (1.41.6 m)

Red-breasted goose
Branta rucollis 212112 in (5355 cm)

Grey francolin
Francolinus pondicerianus
131 2 in (34 cm)

Temmincks tragopan
Tragopan temminckii
25 in (63 cm)


Order Sphenisciformes

Order Gaviiformes

Although some species venture as far

north as the Equator, these distinctive
ightless seabirds are most typically
associated with the very cold
conditions and rich marine food
sources farther south in the
Southern Ocean. Most species
breed in large colonies, some
consisting of hundreds of
thousands of birds. Penguins
have plump bodies and can
weigh between 21 2 and 66 lb
(130 kg). They have short
legs, webbed feet, and wings
that are attened to serve as
ippers. All species stand
upright and waddle on land,
but are superb swimmers,
reaching speeds of up to
9 mph (14 kph). They are
also deep divers, pursuing
sh and krill.

Loons (also called divers) are Northern Hemisphere water

birds, nesting by freshwater, but spending much
time at sea. Their strong legs, set far back on the
body, are ideal for swimming underwater, but
make walking on land impossible.
Arctic loon
Gavia arctica
2312 28 in (6070 cm)

Adelie penguin
Pygoscelis adeliae
1824 in (4661 cm)

Rockhopper penguin
Eudyptes chrysocome
1912 in (50 cm)

Order Podicipediformes
These birds are widespread worldwide and
found on freshwater and at sea on sheltered
inshore waters. They have slim, daggerlike bills,
their feet are broadly lobed, and their legs are
set far back for underwater

King penguin
Aptenodytes patagonicus
35 in (90 cm)

Humboldt penguin
Spheniscus humboldti
23 in (58 cm)

Jackass penguin
Spheniscus demersus
2312 28 in (6070 cm)

Great crested grebe

Podiceps cristatus
1820 in (4651 cm)

Albatrosses, petrels, and shearwaters


Order Procellariiformes

Order Phoenicopteriformes

From tiny storm-petrels to the wandering albatross (which

has the longest wingspan of any bird), albatrosses, shearwaters,
and petrels are all characterized by tubular nostrils. They spend
much time over open sea, but come to land to breed. Smaller
species, effectively unable to walk, visit nests only under cover
of darkness, and even the larger species are weak on land, but
all show great mastery of oceanic conditions.

Flamingos are tropical and temperate waterside birds

that live and breed socially, within very
few, large breeding colonies. Tens
or hundreds of thousands gather
to feed in very restricted
areas with favorable
conditions. They are
long-legged, longnecked, and have
angled bills
used for ltering
food from the water as they
wade, or sometimes swim.

A breeding pair of royal albatrosses (Diomedea
epomophora ) on their nest.

Northern fulmar
Fulmarus glacialis
1712 1912 in (4550 cm)

Corys shearwater
Calonectris diomedea
17 12 22 in (4556 cm)

Greater amingo
Phoenicopterus ruber
443 4 ft (1.21.45 m)

European storm-petrel
Hydrobates pelagicus
51 2 612 in (1417 cm)

Storks, ibises, and herons

Order Ciconiiformes
This is a varied group of waterside birds that includes herons,
bitterns, egrets, storks, ibises, and spoonbills. They are widespread,
but some species are localized, restricted to tiny areas of suitable
habitat such as reed beds. Most are long-legged, long-necked
birds, with daggerlike bills for grasping, not
stabbing, sh. Spoonbills have attened
bills with sensitive, spoon-shaped tips
used to detect food in shallow water.

Black-crowned night-heron
Nycticorax nycticorax
2363 in (5865 cm)

European spoonbill
Platalea leucorodia
3237 in (8093 cm)

Scarlet ibis
Eudocimus ruber
2227 in (5668 cm)
European white stork
Ciconia ciconia
3312 ft (0.951.1 m)

Cattle egret
Bubulcus ibis
1820 in (4550 cm)

Grey heron
Ardea cinerea
3539 in (9098 cm)




Pelicans and relatives

Order Pelecaniformes
A widespread group in tropical and temperate regions, these
birds all share long, angular wings, exible throat pouches, and
forward-facing eyes. All but the frigatebirds have webbing joining
all four toes (unlike ducks or gulls). Many are seabirds, but some
are freshwater species. Colonies sometimes number many
thousands of pairs.

Brown pelican
Pelecanus occidentalis
312 5 ft (11.5 m)

Northern gannet
Morus bassanus
353934 in (89102 cm)

Great cormorant
Phalacrocorax carbo
3139 in (80100 cm)

Balaeniceps rex
31 2 41 2 ft
(1.11.4 m)

Great frigatebird
Fregata minor
3341 in (85105 cm)

Birds of prey

Cranes and rails

Orders Accipitriformes, Cathartiformes, and Falconiformes

Order Gruiformes

A diverse group, birds of prey include birds smaller than a thrush

as well as some of the worlds largest ying birds. Many are
predatory, others scavengers, and some are largely
vegetarian. Most species have muscular legs,
sharp talons, and a sharp, hooked bill.

Characterized by the tall, upright, long-legged,

powerful cranes, these birds are strongly
migratory and social. The group includes birds
as varied as rails and crakes, many of which
are waterside birds; the bustards, which inhabit
various open grassland habitats; and oddities
such as the limpkin and seriemas. The order
is widespread, but many species are secretive,
and some are relatively little-known.

Great bustard
Otis tarda
312 ft (1.1 m)
Turkey vulture
Cathartes aura
2532 in (6481 cm)

Andean condor
Vultur gryphus
4414 ft (1.1.3 m)

Little bustard
Tetrax tetrax
151 2 171 2 in (4045 cm)

Southern caracara
Caracara plancus
191 2 23 in (4959 cm)

Common kestrel
Falco tinnunculus 1215 in (3038 cm)

Rhynochetos jubatus 2112 in (55 cm)

Harriss hawk
Parabuteo unicinctus
30 in (75 cm)
Common moorhen
Gallinula chloropus
121 2 14 in (3235 cm)

Water rail
Rallus aquaticus
81 2 11 in (2228 cm)

Purple swamphen
Porphyrio porphyrio
15191 2 in (3850 cm)

Secretary bird
414 41 2 ft (1.31.4 m)
White-bellied sea eagle Haliaeetus
leucogaster 28 in (70 cm)

Buff-banded rail
Gallirallus philippensis
1113 in (2833 cm)

Egyptian vulture (immature) Neophron

perncopterus 2328 in (5870 cm)

Aramus guarauna 2228 cm (5671 cm)

Common crane
Grus grus
34 ft (0.951.2 m)


Waders, gulls, and auks

Order Charadriiformes
This large and varied order, found worldwide, includes some
of the greatest long-distance migrants of the bird world. The core
groups are wading birds or shorebirds, including plovers and
sandpipers; the more marine gulls and terns; and true seabirds,
the auks, including guillemots and pufns, which breed in
spectacular coastal colonies. Waders show great variation
in size, bill shape, and leg length, according to their various
foods and habitats. They spend most time
close to water, but breed on
tundra, moorland, marshland,
or even farmed habitats.

Masked lapwing
Vanellus miles 1315 in (3338 cm)

Eurasian oystercatcher
Haematopus ostralegus
151 2 1712 in (4045 cm)

Black-necked stilt
Himantopus mexicanus
14 in (35 cm)

Wattled jacana
Jacana jacana
61 2 10 in (1725 cm)
Little ringed plover
Charadrius dubius
51 2 6 in (1415 cm)

Pied avocet
Recurvirostra avosetta
1612 1712 in (4245 cm)

Common gull
Larus canus
15171 2 in (3844 cm)

Eurasian curlew
Numenius arquata
191 2 2312 in (5060 cm)

Long-tailed jaeger
Stercorarius longicaudus
1921 in (4853 cm)

Inca tern
Larosterna inca
151 2 161 2 in (3942 cm)

Atlantic pufn
Fratercula arctica
10 in (25 cm)

Parrots and cockatoos

Order Pteroclidiformes
The sandgrouse are a small group of
terrestrial birds found in warm or hot,
open steppe or semidesert habitats.
These short-legged, stout-bodied, longwinged birds are weak walkers but swift
iers. They often live socially, making
regular morning or evening ights to water.
They can carry water back to their chicks
in their soaked belly feathers.

Pin-tailed sandgrouse
Pterocles alchata
11 in (28 cm)

Order Psittaciformes
Widespread tropical birds, which are
found mostly in scrub, woodland, or forest.
They are often social, and feed on fruit and
seeds. All have short legs, strong feet with
two toes pointing backward, and stout,
hooked bills with bare skin (a cere) at the
base, somewhat like falcons and pigeons.

Nestor notabilis
18 in (46 cm)

White cockatoo
Cacatua alba
18 in (46 cm)

Pigeons and doves

Order Columbiformes
A large group of species, which is found
worldwide on land, except in areas of
extreme cold and desert conditions. The
order comprises pigeons and doves (there
is no clear separation between the two)
and includes two well-known extinct
families, the dodo and solitaires.
Croaking ground
6 in (15 cm)

Speckled pigeon
Columba guinea
131 2 in (33 cm)

Brown lory
Chalcopsitta dulvenbodei
121 2 in (32 cm)

Black-winged lory
Eos cyanogenia
4 in (10 cm)

Mourning dove
Zenaida macroura
91312 in
(2334 cm)
fruit dove
10 in (26 cm)

Red-fronted lorikeet
Charmosyna rubronotata 4 in (10 cm)

St. Vincent parrot

Amazona guildingii 151 2 in (40 cm)


Cuckoos and turacos

Order Cuculiformes
This group is found widely around the world, some cuckoos
being long-distance migrants between tropical and temperate
regions. Turacos are residents of tropical
forests, and the hoatzin lives in South
American forests. Many cuckoos lay
their eggs in the nests of other
species and play no part in
raising their own young.
Cuckoos feed on insects,
especially caterpillars.

Red-crested turaco
Tauraco erythrolophus
151 2 17 in (4043 cm)

This primitive South American hoatzin (Opisthocomus
hoazin ) lives in trees and feeds almost entirely on
leaves. It is 2428 in (6272 cm) in length.

Common cuckoo
Cuculus canorus 121 2 13 in (3233 cm)


Nightjars and frogmouths

Order Strigiformes

Order Caprimulgiformes

Owls and barn owls are mostly crepuscular or nocturnal,

but some feed by day. Most owls have excellent hearing
and eyesight; adapted to almost silent ight, they are able
to pinpoint prey by sound in near darkness or under
snow. They occupy tundra and desert, dense rain forest,
and open scrub. Found worldwide, the barn owl is
among the most widespread of land birds.

Nightjars and frogmouths are found around the world, together

with localized species of Caprimulgiformes, such as the oilbird.
They are active at dawn and dusk or in darkness, usually
hunting for ying insects. Nightjars and frogmouths are
characterized by tiny legs, tiny bills with huge gapes,
large eyes, and highly cryptic plumage patterns. By
day they rely on their camouage to hide against
bark, in foliage, or on the ground.

European nightjar
Caprimulgus europaeus 1011 in (2628 cm)

Eagle owl
Bubo bubo
2329 in (5973 cm)

Spectacled owl
Pulsatrix perspicillata
172012 in (4352 cm)

Tawny owl
Strix aluco
1412 151 2 in (3739 cm)

Tawny frogmouth
Podargus strigoides
1312 21 in (3453 cm)

Common potoo
Nyctibius griseus
1315 in (3338cm)

and swifts
Order Apodiformes
Swifts are the worlds most aerial
birds, some not coming to land for
two or three years, and then only
to nest. They catch insects, despite
having tiny bills. The hummingbirds
eat nectar, having highly adapted
bill shapes. The worlds most
maneuverable birds, they hover,
dash forward, or can even y
Pallid swift
Apus pallidus
backward. This group includes
61 2 7 in
the worlds smallest birds.

Great grey owl

Strix nebulosa
2333 in (5985 cm)

Andean hillstar
estella 56 in
(1315 cm)

(1618 cm)

Eurasian pygmy owl

Glaucidium passerinum
67 1 2 in (1519 cm)

Order Coliiformes
Just six species, all found in tropical
Africa, make up this order. They are
characteristic of open bush in the great
savanna plains. Mousebirds have small,
decurved bills, short but strong feet,
and long, slender tail feathers. They
are highly social birds, feeding close
together. This is one of only two
orders of birds exclusively native
to Africa. The other is the ostrich.

Ural owl
Strix uralensis
2324 in (5862 cm)

Northern hawk-owl
Surnia ulula
1416 in (3641 cm)

Barn owl
Tyto alba
1112 17 1 2 in (2944 cm)

Blue-naped mousebird
Urocolius macrourus
1314 in (3336 cm)

Speckled mousebird
Colius striatus
121512 in (3040 cm)

Kingfishers and relatives

Order Trogoniformes

Order Coraciiformes

Despite being found in the widely

separated tropical regions of Africa, the
Americas, and Indo-China, all trogons
are remarkably similar in appearance.
They are essentially green above, often
barred black-and-white on the tail, and
bright red, pink, orange, or yellow below.
They have short, stout bills, and short legs.
Most species prefer forest or woodland
habitats and feed on insects.

Kingshers, motmots, bee-eaters, rollers, hoopoes,

and hornbills as a group are represented almost worldwide,
but only kingshers are really widespread. Few kingshers
actually eat sh, many catching insects or reptiles in
woodland, as do the Australasian kookaburras. Bee-eaters
take insects on the wing; rollers drop from perches to the
ground to catch their prey; hoopoes and some hornbills
feed on the ground. Kingshers range from tiny to
medium sized, whereas ground hornbills are
huge birds. These species all nest in holes of
various kinds, either tunneled into earth or
ready-made in tree cavities.

Resplendent quetzal
Pharomachrus mocinno
14151 2 in (3540 cm)

Violaceous trogon
Trogon violaceus
910 in (2326 cm)



European roller
Coracias garrulus
12 in (30 cm)

Woodpeckers and toucans

Order Piciformes
Puffbirds, barbets, honeyguides, toucans,
and woodpeckers form a group with zygodactyl
feetmeaning that they have two toes facing
forward and two toes facing backward.
They have strong bills and often rather
bold, noisy behavior. Honeyguides
lead other animals to bees nests
and benet from the other animals
breaking into the hives. Toucans
are characterized by remarkable,
colorful, long but lightweight
bills. Woodpeckers use their
tails as props to support them
against upright branches, and
drum against branches with
their bills, using this instrumental
communication in place of
conventional song. Only the
woodpeckers are widespread
on several continents.

Toco toucan
Ramphastos toco
21231 2 in (5360 cm)

Laughing kookaburra
Dacelo novaeguineae
161 2 in (42 cm)

Pied kingsher
Ceryle rudis
10 in (25 cm)

Common kingsher
Alcedo atthis 61 2 in (16 cm)

Blue-crowned motmot
Momotus momota 1812 in (47 cm)
Jamaican tody
Todus todus
412 in (11 cm)

Common hoopoe
Upupa epops
1011 in (2628 cm)
DArnauds barbet
Trachyphonus darnaudii
8 in (20 cm)

Northern wryneck
Jynx torquilla
61 2 in (16 cm)

Middle spotted woodpecker

Dendrocopos medius
712 81 2 in (1922 cm)

Greater honeyguide
Indicator indicator
8 in (20 cm)

Galbula rucauda
10 in (25 cm)

Southern yellow-billed hornbill

Tockus leucomelas
191 2 231 2 in (5060 cm)

Great hornbill
Buceros bicornis
5 ft (1.5 m)

European bee-eater
Merops apiaster
12 in (30 cm)


Order Passeriformes
By far the largest order of birds, all
passerines have four unwebbed toes with
three pointing forward, one backward.
They are otherwise hugely variable in size,
form, color, behavior, and habitat. Known
as the perching birds, they form two varied
suborders. The smaller group, the suboscine passeriformes, has 12 families,
and the larger, the oscines, often called
songbirds, has 70 families. Some songbirds,
such as the crows, are not very musical,
while others, such as the Australasian
lyrebirds, Old and New World thrushes, and
various chats, including the nightingale, are
among the worlds nest songsters.

Acanthisitta chloris
31 2 in (3 cm)

Green broadbill
Calyptomena viridis
8 in (20 cm)

African pitta
Pitta angolensis
8 in (20 cm)

Barred antshrike
Thamnophilus doliatus
61 2 in (16 cm)
Three-wattled bellbird
Procnias tricarunculatus
1012 in (2530 cm)

Blue manakin
Chiroxiphia caudata
5 in (13 cm)

Red-backed fairy-wren
Malurus melanocephalus
45 in (1013 cm)

Oxyruncus cristatus
61 2 in (17 cm)

Buff-fronted foliage-gleaner
Philydor rufum. 7 1 2 in (19 cm)

Blue-faced honeyeater
Entomyzon cyanotis
12 in (31 cm)

Lesser grey shrike

Lanius minor 71 2 8 in (1920 cm)

Eastern kingbird
Tyrannus tyrannus
8 in (20 cm)

Wedge-billed woodcreeper
Glyphorynchus spirurus
51 2 in (14 cm)

Despite its extraordinary appearance, the lyrebird
(Menura novaehollandiae ) is elusive. It is known for
its remarkable mimicry of natural and artificial sounds.

Brown thornbill
Acanthiza pusilla
4 in (10 cm)

Tawny-crowned greenlet
Hylophilus ochraceiceps 41 2 in (11.5 cm)

Grey-chinned minivet
Pericrocotus solaris
71 2 in (19 cm)

Eastern black-headed oriole

Oriolus larvatus 81 2 in (22 cm)


Grallina cyanoleuca
111 2 in (29 cm)

Scarlet robin
Petroica boodang
434 512 in (1214 cm)

Dulus dominicus
7 in (18 cm)

Carrion crow
Corvus corone
201 2 in (52 cm)

Blue jay
Cyanocitta cristata
12 in (30 cm)

White-necked picathartes
Picathartes gymnocephalus
151 2 191 2 in (3950 cm)

Blue tit
Parus caeruleus 41 2 in (12 cm)

Bank swallow
Riparia riparia
41 2 in (12 cm)

Dartford warbler
Sylvia undata
41 2 5 in (1213 cm)

Red-whiskered bulbul
Pycnonotus jocosus
8 in (20 cm)

Bearded tit
Panurus biarmicus
41 2 6 in (1215 cm)

Bohemian waxwing
Bombycilla garrulus 612 in (17 cm)

Asian fairy-bluebird
Irena puella
1012 in (27 cm)

Cape white-eye
Zosterops pallidus 41 2 in (11 cm)

Greater bird-of-paradise
Paradisaea apoda
14 in (35 cm)

Eurasian skylark
Alauda arvensis
77 1 2 in (1819 cm)

Red-billed leiothrix
Leiothrix lutea
6 in (15 cm)

Winter Wren
Troglodytes troglodytes
31 2 4 in (910 cm)


Common starling
Sturnus vulgaris
81 2 in (21 cm)

Eurasian nuthatch
Sitta europaea
48 in (1020 cm)

Song thrush
Turdus philomelos
9 in (23 cm)

House sparrow
Passer domesticus
51 2 in (14 cm)

Northern mockingbird
Mimus polyglottos
911 in (2328 cm)

Eurasian treecreeper
Certhia familiaris
5 in (12.5 cm)

Rufous-gorgetted ycatcher
Ficedula strophiata
51 2 in (14 cm)

Scarlet-chested sunbird
Chalcomitra senegalensis 6 in (15 cm)

Java sparrow
Lonchura oryzivora
51 2 6 in (1415 cm)

Prunella modularis
51 2 in (14 cm)

Bullocks oriole
Icterus bullockii
81 2 in (21 cm)

Hooded warbler
Wilsonia citrina 51 2 in (14 cm)

Eurasian bullnch
Pyrrhula pyrrhula
6 in (15 cm)

Snow bunting
Piectrophenax nivalis
61 2 in (16 cm)

Richards pipit
Anthus richardi
61 2 8 in (1720 cm)

Green honeycreeper
Chlorophanes spiza 51 2 in (14 cm)

Northern cardinal
Cardinalis cardinalis 81 2 in (22 cm)


Related to Asian parrotbills rather than
true tits, the bearded tit (Panurus biarmicus )
is usually confined to reed beds. When
numbers are high in fall, some disperse
in search of new sites and may be seen
in cattails or other marsh plants.





Mammalia Characteristics of this group include a lower

jaw comprising a single bone, and skin that contains glands
(including mammary glands) and is usually covered with hair.

Theria Mammals that produce live young, at various

stages of development. This group includes both the
placental and marsupial (pouched) mammals.

Placentalia Live-bearing mammals with young that develop

in the uterus attached to a placenta which allows nutrients
and waste to pass between a mother and her offspring.

Page 36



Species 31


Species 87

Species 21



Prototheria The only mammals that reproduce by laying

soft-shelled eggs. Their body temperature is maintained
at a lower level than that of most other mammals.

Marsupialia Live-bearing mammals that have a pouch

(marsupium) in which the young are nurtured on milk to
complete their development.




Species 143

Species 80

Species 5

Laurasiatheria The most diverse group of placental
mammals, ranging from whales to bats, cats
to rhionoceroses.



Species 1,116

Species 452

Species 12

Species 14



Species 67

Species 1


Species 16


Species 35

Carnivora Mainly meat-eating mammals with

well-developed canine teeth and powerful cheek teeth
with a scissorlike action, called carnassials.

Species 19

Species 8

Species 35

Species 7



Species 1




Species 38

Species 4

Species 4

Species 143


Species 36



Species 8

Species 51

Species 19

Cetartiodactyla Laurasiatherians with a uniquely shaped
hock bone, known as the astragulus or talus, including the
terrestrial ancestors of modern whales.


Species 2

Species 84

Species 2

Mammal groups
The primitive egg-laying mammals were the rst group to
diverge, followed by the marsupials. The placental mammals
divided into three broad groups, the laurasiatherians,
afrotherians, and euarchontoglires, plus the anteaters and
relatives, before evolving into the diverse array of forms

and sizes we see today. Although many of the traditional

mammal groups, such as primates and carnivores, appear
on this diagram, others are less familiar. These include the
even-toed hooved mammals and whales, which have been
combined quite recently using genetic and fossil evidence.



Afrotheria A group representing an ancient radiation
of African mammals. Now diversied, they bear little
outward resemblance to one another.

Species 4


Species 3

Species 5


Species 92


Species 1

Species 51


Species 1,518

Species 15



Species 2

Species 40



Euarchontoglires A group of placental mammals

that combines rodents and rabbits with primates, tree
shrews, and colugos.

Rodentia Gnawing mammals with a single pair of incisor

teeth in the upper and lower jaws that are open-rooted and
grow throughout life.



Species 8

Species 4


Species 235

Species 184

Species 278


Species 20

Perissodactyla Plant-eating mammals with an odd

number of weight-bearing toes. Cellulose-digesting
bacteria are housed in the hind gut.

Species 2

Species 5



Species 37

Species 7

Species 5

Species 17

Primates Mammals with grasping extremities,
binocular vision, and large brains.

Species 128

Species 153


fat. Milk-producing mammary glands are also

found only in mammals. The milk provides
nourishment to offspring, removing the need to
expend energy on foraging, and initially contains
antibodies, which protect infants from disease.

Despite their relatively late appearance in the fossil record, mammals have
diversied into an astonishing array of shapes and sizes. They are most
diverse and widespread on land, but have also colonized air and water.
About 5,000 species of mammal have been
described, ranging from the egg-laying duckbilled platypus to humans with their sophisticated
brain and hand dexterity, but there are similarities
that unite them all in the class Mammalia.
Mammals uniquely possess fur, which protects
delicate skin, provides camouage, and helps

Key characters

insulate the body. Mammals also have skin

glands, among which are sebaceous glands
that allow sweating. Mammals are endothermic,
which means that they maintain a constant
body temperatureoften above that of their
surroundingsby producing heat through
metabolic processes, such as breaking down

The only characters that can identify all mammals,

whether extant (living) or fossil, are skeletal
among these are the possession of a single
bone in the lower jaw (dentary) and having
three bones (incus, stapes, and malleus) in the
middle ear. Any one of these features identies
an animal as a mammal and separates it from all
other living things.

Order Monotremata
All other mammals produce live young,
but monotremes lay soft-shelled eggs,
which hatch after a short incubation
period. Monotremes have only a single
posterior opening, called the cloaca, into
which the urinary, alimentary, and
reproductive systems open.

Duck-billed platypus
Ornithorhynchus anatinus
1624 in (4060 cm)

Short-beaked echidna
Tachyglossus aculeatus 121712 in (3045 cm)

Supercohort Marsupialia
These are the so-called pouched mammals, although not all of
the species have one. The offspring are born after a very limited
gestation period, and then make their way to the pouch. Born
in almost embryonic form, the young attach to a nipple and
suckle milk while completing their development.
Marsupialia is divided into seven orders, including:
American opossums (Didelphimorphia);
bandicoots and bilbies (Paramelemorphia);
and koalas, kangaroos, wombats,
and possums (Diprotodontia).

Virginia opossum
Didelphis virginiana
1520 in (3850 cm)

Feather-tailed possum
Distoechurus pennatus
412 512 in (10.513.5 cm)
Greater glider
Petauroides volans
1419 in (3548 cm)

Rufus bettong
Aepyprymnus rufescens 1412
2012 in (3752 cm)

Common brushtailed possum

Trichosurus vulpecula
1423 in (3558 cm)

Red-necked wallaby
Macropus rufogriseus
2841 in (70105 cm)

Common wombat
Vombatus ursinus 2847 in (70120 cm)

Phascolarctos cinereus
2631 in (6578 cm)
Setonix brachyurus
162112 in (4054 cm)

Tasmanian devil
Sarcophilus harrisii
2842 in (70110 cm)

Red kangaroo
Macropus rufus
314 514 ft (11.6 m)

This tiny Australian marsupial (Tarsipes rostratus )
is only 21 2 31 2 in (6.59 cm) long, and gathers
pollen and nectar (honey) from flowers.

Elephant shrews



Order Tubulidentata

Order Hyracoidea

Order Afrosoricida

Order Macroscelidea

This recently recognized group is chiey

supported by genetic evidence, but there
are some features of the dentition that are
characteristic. Afrosoricids have only one
major cusp on each of the upper molars;
other placental mammals have several.

Elephant shrews, or sengis, are

characterized by a long, highly mobile
snout, an enlarged cecum (a pouch
connected to the large intestine) in the
hind gut, and cheek teeth
that can grow
throughout life
since they
have open

This monospecic order comprises

only the long-eared, long-snouted
aardvark. Unlike most other mammals,
its teeth lack an outer coating of enamel
and are covered instead with cementum,
the same tissue that coats tooth roots.

These species have a unique eye

structure. A part of the iris projects
over the pupil. This has the effect of
reducing the amount of overhead light
entering the eye.

Common tenrec
Tenrec ecaudatus 10151 2 in (2539 cm)

Rufous elephant shrew

Elephantulus rufescens 412 5 in (1212.5 cm)

Order Proboscidea
An elephant can be easily recognized by its large size,
columnar legs, and long trunk. The incisor teeth are modied
into continually growing tusks of dentine, while the molar
teethof which there are six in each half of the upper and lower
jawserupt at the back of the jaw and migrate forward in a
conveyer-belt-like fashion over about 60 years. The large head
is made lighter by the presence of air pockets in the skull.

Rock hyrax
Procavia capensis
2123 in (3058 cm)

Orycteropus afer
514 ft (1.6 m)

Dugongs and
Order Sirenia
These large, slow-moving creatures
are streamlined for their aquatic lifestyle.
Sirenian forelimbs are modied into
ippers, while the hindlimbs have been
entirely lost. Instead there is a horizontal,
spatula-like tail. Manatees have only six
neck vertebrae compared to the seven
found in most other mammals. Entirely
herbivorous, sirenians have unusually long
guts, with bacteria for the digestion of
cellulose. This bacteria is housed in the
cecum in the hind portion of the gut, as
it is in horses. They are the only marine
mammals that feed purely on plants.
Cheek tooth replacement is similar to
that of the elephant.

West Indian manatee

Trichechus manatus
814 1434 ft (2.54.5 m)

Dugong dugon
814 13 ft (2.54 m)

Anteaters and relatives

Orders Pilosa, Cingulata

African elephant
Loxodonta africana
131612 ft (45 m)

The order Pilosa comprises anteaters and sloths, and

the order Cingulata contains the armadillos. Both groups
are unique in having a double vena cava vein in the
lower part of the body and additional moving joints, or
articulationscalled xenarthraleson the lower lumbar
vertebrae. The two groups can easily be separated
because the armadillos have an armored upper body
whereas anteaters and sloths have fur.

Dasypus sp.
12151 2 in (3040 cm)

Asian elephant
Elephas maximus 1112 ft (3.5 m)

Giant anteater
Myrmecophaga tridactyla
314 612 ft (12 m)

Two-toed sloth
Choloepus didactylus
1834 in (4686 cm)

Southern tamandua
Tamandua tetradactyla 2135 in (5388 cm)

Silky anteater
Cyclopes didactylus
61 2 81 2 in (1621 cm)


Tenrecs and
golden moles


Order Rodentia
These are the gnawing mammals. Most are small, but all have
characteristic teeth. Rodents have a single pair of continually
growing incisor teeth in the upper and lower jaws that are sharpened
as the lower ones sheer against the inside of the upper ones. Canine
teeth are absent, leaving a gap called the diastema. Many species
have broad, ridged cheek teeth, which are effective in grinding up
vegetation, but some rodents are carnivorous. The cavylike
rodents can be separated from the other rodents on the basis
of their jaw musculature and longer gestation periods that lead
to the birth of smaller numbers of well-developed offspring.

Canadian beaver
Castor canadensis
2935 in (7488 cm)

Gray squirrel
Sciurus carolinensis
912 in (2330 cm)

Alpine marmot
Marmota marmota
1912 211 2 in (5055 cm)

Pedetes capensis
1012 1512 in
(2740 cm )

Spiny mouse
Acomys cilicius 612 7 in (1718 cm)

Brown rat
Rattus norvegicus
812 111 2 in (2129 cm)

Forest dormouse
Dryomys nitedula
314 5 in (813 cm)

Striped grass mouse

Lemniscomys striatus
451 2 in (1014 cm)

Malagasy giant rat

Hypogeomys antimena 1214 in (3035 cm)

Bank vole
Clethrionomys glareolus 314 412 in (811 cm)

Red vole
Clethrionomys rutilus 314 412 in (811 cm)

Four-toed jerboa
Allactaga tetradactyla
441 2 in (1012 cm)

North American porcupine

Erethizon dorsatum
2631 in (6580 cm)

Long-tailed chinchilla
Chinchilla lanigera 812 9 in (2223 cm)

Cape porcupine
Hystrix africaeaustralis 1912 in (50 cm)

Naked mole-rat
Heterocephalus glaber
512 7 in (1418 cm)

Hydrochoerus hydrochaeris
312 414 ft (1.11.3 m)

Rabbits, hares, and pikas

Order Lagomorpha
These are herbivorous mammals that gain maximum
nutrients from food by ingesting it twice, a behavior called
coprophagy. Rabbits and pikas also have very distinctive
upper incisor teeth, with a pair of small, nonfunctional,
peglike incisors behind the functional pair. Like rodents, they
lack canines, which creates a large gap through which the
lips can be drawn to prevent debris from entering the mouth
while feeding.

Hispid hare
Caprolagus hispidus
151912 in (3850 cm)

Oryctolagus cuniculus
141912 in (3550 cm)

North American pika

Ochotona princeps 612 812 in (1622 cm)


Order Primates
Members of this order are characterized by an opposable thumb
or toe and the ability to rotate the lower arm bones (radius and
ulna) around one another. These features allow primates to grasp
and manipulate objects. The order is generally subdivided. The
lemurs, galagos, and lorises form the strepsirhines, which depend
more on their sense of smell than do monkeys and apes. They
generally have longer snouts, a glandular (and therefore moist) and
naked nasal region, and comma-shaped nostrils. The tarsiers,
marmosets, monkeys, and apes form the second group, the
haplorhines, which have dry noses with ovate nostrils.

Gray woolly monkey

Legothrix cana
191 2 26 in (5065 cm)

Ring-tailed lemur
Lemur catta
1512 18 in (3946 cm)

Slender loris
Loris tardigradus
612-10 in (1726 cm)

Red howler monkey

Alouatta seniculus
2025 in (5163 cm)

Senegal bushbaby
Galago senegalensis 661 2 in (1517 cm)

Bolivian squirrel monkey

Saimiri boliviensis
1012 1212 in (2732 cm)

Golden lion tamarin

Leontopithicus rosalia
810 in (2025 cm)

Patas monkey
Erythrocebus patas
2312 35 in (6088 cm)

Black and white colobus monkey

Colobus guereza
2012 2212 in (5257 cm)

Theropithecus gelada 2829 in (7074 cm)

Proboscis monkey
Nasalis larvatus 2930 in (7376 cm)

Lar gibbon
Hylobates lar 1612 23 in (4259 cm)

Western gorilla
Gorilla gorilla
414 614 ft (1.31.9 m)

Mandrillus sphinx
2532 in (6381 cm)

De Brazzas monkey
Cercopithecus neglectus
191 2 23 in (5059 cm)

Pan troglodytes
212 3 ft (0.751 m)

Hylobates syndactylus
35 in (90 cm)

Pongo pygmaeus 33 ft 3 in (0.91 m)


Tree shrews

Flying lemurs


Order Scandentia

Order Dermoptera

Order Eulipotyphla

These rather squirrel-like mammals have

slender bodies and long tails. In the past
they have been classied as insectivores
and as primates on the basis of characters
they have in common with these orders.
However, genetic evidence now suggests
that tree shrews are an ancient group with
an independent evolutionary history.

Flying lemurs, or colugos, are cat-sized,

arboreal mammals. Their distinctive feature
is the large gliding membrane, called a
patagium, which stretches from the neck
to the front and hind digits and onto the
very tip of the tail.

These insect-eating animals

have a small brain, which lacks
the copious infolding found in
other mammals. In common with
monotremes and marsupials, they
have a cloaca, but unlike them the
young develop longer in the womb.

European mole
Talpa europaea
41 2 61 2in (1116cm)

Lesser tree shrew

Tupaia minor
412 512 in (11.513.5 cm)

Malaysian ying lemur

Cynocephalus variegatus
131612 in (3342 cm)

European hedgehog
Erinaceus europaeus
812 in (2030 cm)

Common shrew
Sorex araneus
214 312 in (5.59 cm)

Lesser mouse-tailed bat

Rhinopoma hardwickei
214 234 in
(5.57 cm)

Order Chiroptera
Bats are the only mammals that are capable of powered ight.
Their forelimbs have very long nger bones that support the ight
membrane. Some bats, known as ying foxes, feed only on fruit
and nectar. They do not echolocate because they have
excellent vision and sense of smell. Most bats eat insects,
which they locate by emitting ultrasound pulses and
listening to the returning echoes. To pinpoint the
sounds these bats may have large ears with an
additional projection called a tragus.

Lesser horseshoe bat

Rhinolophus hipposideros
11 2 13 4 in (3.54.5 cm)
Rodrigues ying fox
Pteropus rodricensis
14 in (35 cm)

Noctule bat
Nyctalus noctula
214 314 in (68 cm)
Sebas short-tailed bat
Carollia perspicillata
2212 in (56.5 cm)

Daubentons bat
Myotis daubentonii
134 214 in (4.55.5 cm)

Order Pholidota
Distinctive, overlapping body scales
provide pangolins with a unique protective
armor. Since they feed primarily on ants
and termites, they lack teeth and the lower
jaw is much reduced. With no teeth to grind
up their food, pangolins have a tough,
muscular stomach that does this instead.

Common vampire bat

Desmodus rotundus 234 31 2 in (79 cm)

Whiskered bat
Myotis mystacina
11 2 2 in (3.55 cm)

Cape pangolin
Manis temminckii
1628 in (4070 cm)

Brown long-eared bat

Plecotus auritus
112 2 in (45 cm)

Parti-colored bat
Vespertilio murinus
2212 in (56.5 cm)


Order Carnivora
A predominantly meat-eating group, carnivora includes
terrestrial and aquatic species. The terrestrial carnivores are
characterized by their large canine teeth and specialized
carnassialslast upper premolar and rst lower molarused
for slicing through hide, meat, and bone. In seals and walruses
the limbs are modied into ippers and the teeth are
less specialized because these animals feed on
sh and invertebrates that are swallowed whole.
Since meat is relatively easy to digest, the stomach is simple and
the intestine short. In all carnivores the clavicle bone is lost, or very
reduced, and three of the small bones in the wristthe scaphoid,
centrale, and lunar boneshave fused to form a scapholunar bone.

Least weasel
Mustela nivalis
612 912 in (16.524 cm)

California sea lion

Zalophus californianus
7 ft (2.1 m)

Red panda
Ailurus fulgens
2025 in (5064 cm)

Brown bear
Ursus arctos
512 914 ft (1.72.8 m)

Acinonyx jubatus
312 5 ft (1.11.5 m)

Caracal caracal
2436 in (6091 cm)

Odobenus rosmarus
934 1134 ft (33.6 m)

Sand fox
Vulpes rueppellii
16201 2 in (4052 cm)

Panthera tigris
612 1214 ft (23.7 m)

Panthera leo
51 2 814 ft (1.72.5 m)

Banded mongoose
Mungos mungo 211 2 24 in (5560 cm)

Suricata suricatta 1014 in (2535 cm)

Eurasian badger
Meles meles
2235 in (5690 cm)

Procyon lotor
2436 in (60100 cm)

Polar bear
Ursus maritimus
up to 814 ft (2.5 m)

Grey wolf
Canis lupus 11.5m (314 5ft)


Horses, tapirs,
and rhinoceroses
Order Perissodactyla
Odd-toed ungulates are so-called
because they bear their weight on either
one (equids) or three (rhinoceroses and
tapirs) toes. Being plant eaters, they need
cellulose-digesting bacteria in their guts to
break down plant cell walls. These bacteria
are housed in the cecuma blind-ended
sac leading off the small intestinehence
they are referred to as hindgut digesters.

African wild ass

Equus asinus
61 2 712 ft (22.3 m)

Brazilian tapir
Tapirus terrestris
512 612 ft (1.72 m)

White rhinoceros
Ceratotherium simum
1214 13 ft (3.74 m)

Terrestrial even-toed
Order Cetartiodactyla (part)
The order Cetartiodactyla comprises
what used to be recognized as two separate
orders, Artiodactyla (terrestrial even-toed
ungulates) and Cetacea (whales, dolphins,
and porpoisessee opposite). The recent
recognition that these two seemingly
different groups belong together is based
mainly on genetic evidence, but backed up
by some fossil evidence. The land-dwelling
species are known as even-toed ungulates,
because they bear their weight on two toes,
each enclosed in a hoof of keratin. Many
species ruminatethat is they regurgitate
and further chew their plant food. They are
also called foregut digesters, as they have
digestive bacteria in their rumen, the rst of
the four chambers of the stomach. A major
food source for the larger carnivores, these
ungulates have laterally situated eyes for
all-round vision and most have elongated
lower legs that enable them to be energy
efcient when running for long periods.

Phacochoerus africanus
35 ft (0.91.5 m)

Collared peccary
Tayassu tajacu 3039 in (75100 cm)

Hippopotamus amphibius
1034 1112 ft (3.33.5 m)

Camelus dromedarius
714 1114 ft (2.23.4 m)

Lama glama
37 ft (0.92.1 m)

South African giraffe

Giraffa giraffe
12121512 ft (3.84.7 m)

Alpine musk deer

Moschus chrysogaster
2839 in (70100 cm)

Red deer
Cervus elaphus
5612 ft (1.52 m)

American bison
Bison bison
71112 ft (2.13.5

Okapia johnstoni 612 714 ft (22.2 m)

Scimitar-horned oryx
Oryx dammah 412 734 ft (1.42.4 m)

Ovis musimon
312 414 ft (1.11.3 m)


Whales, dolphins, and porpoises

Order Cetartiodactyla (part)
These mammals are perfectly adapted to an aquatic lifestyle,
having streamlined bodies with ippers for forelimbs and a large
tail uke instead of legs. The skeleton is much reduced, serving
mainly for muscle attachment, and the neck vertebrae may be
fused. The whales and dolphins can be divided into two groups
those with teeth and those without. The giant baleen whales lter
feed, trapping food particles in two huge baleen plates that hang
down from the sides of the upper jaw. Most toothed whales are
smaller and some can echolocate to nd food.

Common dolphin
Delphinus delphis
712 812 ft (2.32.6 m)

Ganges river dolphin

Platanista gangetica
7814 ft (2.12.5 m)

Pontoporia blainvillei
414 512 ft (1.31.7 m)

Killer whale
Orcinus orca up to 26 ft (8 m)

Commersons dolphin
Cephalorhynchus commersonii
41 2 51 2 ft (1.41.7 m)

Harbour porpoise
Phocoena phocoena 41 2 612 ft (1.42 m)

Dalls porpoise
Phocoenoides dalli
71/4 73/4 ft (2.22.4 m)

Monodon monoceros
131434 ft (44.5 m)

Sperm whale
Physeter catodon
3666 ft (1120 m)

Pygmy sperm whale

Kogia breviceps 83/4 93/4 ft (2.73 m)

Cuviers beaked whale

Ziphius cavirostris
2325 ft (77.5 m)

Northern bottlenose whale

Hyperoodon ampullatus
2033 ft (610 m)

Gray whale
Eschrichtius robustus
4349 ft (1315 m)

Humpback whale
Megaptera novaeangliae
4659 ft (1418 m)

Northern right whale

Eubalaena glacialis
4356 ft (1317 m)

Brydes whale
Balaenoptera edeni
305112 ft (915.5 m)

Pygmy right whale

Caperea marginata
1821 ft (5.56.5 m)


Up to 25 percent of a crabs total weight may be
its shell. This is not simply an outer protective
casing, it is also its skeletona complex set
of supporting structures linked at flexible joints,
enclosing and moved by intricate sets of muscles.

Most members of the animal kingdom possess some kind of strong body
framework, and pulling devices with which to move it. Although the principles
and detailed structure of muscles are virtually constant across all major groups
of animals, types of skeletons show huge variety in design and construction.

Movement and structure

Muscle tissue is the primary means of movement in almost all animals,
except for sponges, allowing an animal to implement its behaviors and
actions. Muscle is often the most plentiful tissue in the body, and it has one
basic function: to shorten, or contract. As it does so, it moves parts of the
skeleton or other body structures. Muscles power not only the movements
visible on the outside, they are also the basis of internal activities, such as
the pumping of the heart. For invertebrates with a hard outer framework,
or exoskeleton, most muscles are positioned on
the inside and attached to the skeletons inner walls.
In vertebrates, the situation is reversed, with the
muscles attaching to the outside of the endoskeleton.
In soft-bodied animals like worms, muscle tissue
forms its own uid-pressurized water skeleton.
tendon attaches
muscle to bone
main part of muscle (also
called belly of muscle)

This scanning electron
micrograph (SEM) of a
skeletal muscle shows
striations (bands) of
muscle fibers.

biceps relaxes
triceps contracts

biceps contracts
triceps relaxes


Two muscles in a vertebrate forelimb
demonstrate a simple antagonistic
system, where each muscle opposes
the pull of the other. If both muscles
exert tension, the joint can be held
steady in any position through its range.

teams. In the simplest arrangement, one muscle pulls a skeletal element

or body part one way, while its opposing partner on the other side relaxes.
To move the part the other way, the opposing muscle contracts while the
rst muscle relaxes. However, this two-way action is given greater range
since there are usually several muscles involved, attached to the skeleton
at varying places and angles, to give differing lines of pull. This allows for
movement with close control in several directions.


perimysium (sheath)
epimysium (sheath of connective
tissue covering entire muscle)


A vertebrate muscle is attached to the skeleton,
usually at each end, and consists of bundles
(fascicles) of muscle fibers (myofibers). Each
fiber is composed of perhaps thousands of
thinner muscle fibrils (myofibrils).

extensor muscle

Muscle arrangement
A typical mammal has more than 600 individual muscles, while some
insects have three times that number. The inner structure of a typical
muscle is based on cellular components known as muscle bers or
myobers. In larger animals, some myobers exceed 3 ft 3 in (1 m) in
length, yet each is thinner than a human hair. Contraction occurs when
bundles of overlapping protein laments in the bers slide past each other.
Because muscles only contract and pull, they are arranged in opposing
In arthropods the muscle structure
can be quite similar to that of
vertebrates, but the arrangement
differs in that the muscles connect
to the inner walls of the exoskeleton.

Most animals have a skeleton or similar framework made up of numerous

parts, which move in relation to each other at joints. In arthropods, the
joints are relatively simple thinnings of the exoskeleton. The cuticle
forms a exible articular membrane, but the harder, rigid layers of
chitin or mineralization are almost absent. In most
vertebrates, the bones of the skeleton are covered
in their joints by cartilage, which reduces friction and
wear. The joint is enclosed in a joint capsule, inside
which there is a lubricating liquid, synovial uid, to
further reduce rubbing. Strong, stretchy ligaments
are attached to the bones and allow the joint to ex.

muscle connecting
muscle for changing
shape of segment
muscle for moving leg

flexor muscle

The frogs tiny joints
have a greater range
of movement than
invertebrate joints.
bone marrow

bearing surface
or pivot

joint capsule


The exoskeleton is flexible at an invertebrate joint,
and the shape of the bearing surfaces usually
allows movement in only one plane or direction. In
a vertebrate joint, a thin layer of synovial fluid helps
prevent friction between the connecting bones.

synovial fluid





Water skeletons
Fluid-based skeletons are found in a huge variety of invertebrates,
especially worms and similar soft-bodied animals. Far from being
soft and oppy, many of these can become rigid and resistant when
muscular action compresses uid inside an animals body.


Most nematode worms, such as this parasitic
roundworm, have only longitudinal muscles
in the body wall. As these contract on each
side alternately, they produce characteristic
C- or S-like thrashing movements.

An animals hydrostatic skeleton, or hydroskeleton, employs similar principles

to hydraulic systems in machinery. In animals, a liquid in some sort of chamber
or container is pressurized by contraction of the muscular walls around it.
The liquid is the internal body uid of the animal. Its compression makes the
structure become hard and rigid, forming a rm skeletal unit. The skeleton
then gives support and protection to the animals body parts. In simpler
animals the whole body covering acts as the hydroskeleton. In more
complex ones, especially segmented (annelid) worms, the pressure
can be limited to selected body compartments.
Muscle fibers compress fluid within a
jellyfishs main body, causing contractions
to run from the center of the bell to its
edges. This produces a pulsing
movement for swimming.

edge of gastric pouch

ring canal




alimentary canal
circular muscle

longitudinal muscle



Each worm segment has a set of circular
muscles around it. Some longitudinal muscles
span a segment, while others run along many
segments. This allows the worm to stretch
some parts, while shortening others.


edge of

Octopus suckers are able

to grasp objects rmly.

Sea cucumber tentacles rely

on internal pressure.


Coral polyps use muscle bers in

the stalks to lean in any direction.

Anemone tentacles bend to the

side on which the muscle contracts.

In a hydroskeleton, changes in muscle tone and the

arrangement of muscles can alter the pressure within
its chambers, thereby changing the skeletons shape,
and causing the structure to become rigid. This
provides a stiff base against which movements can
occur, and also produces the movements themselves.
In many worms, for example, there are two layers
of muscle in the outer body wall: ringlike circulars;
and strap-shaped, longitudinal (lengthwise) muscles.
Contraction of circular muscles squeezes the body,
making it longer and thinner; longitudinal muscle
contraction makes it shorter and fatter; and when
both sets contract, the body
becomes tense, stiff, and rigid.
Apart from whole body
Other combinations of contractions
movements, hydrostatic
and hydraulic principles
permit further movements. If the
can also be used to
longitudinal muscles along one
move smaller body
side shorten, the body curves
parts or individual
appendages. Such
to that side. Using such a system
movements allow
of contractions, burrowing
animals to perform tasks
earthworms are able to push
such as self defense
and capturing prey.
their way between tightly packed
soil particles with great force.

83 Horny skeleTons

Horny skeletons
Horny describes substances that are strong, tough, and resilient, yet which
can also be slightly compressed and flexed. In arthropods, the main horny
substance is chitin. This forms the basis of the outer body covering or cuticle,
which is the main component of the animals exoskeleton.

Tough exoskeleton
Chitin is light, strong, translucent, and pliable, and
has been compared to plastic. The substance is
chemically a polysaccharide (carbohydrate), consisting
of glucoselike sugar units. In most land arthropods
chitin is accompanied by various proteins, the
molecules of which may take many shapes, including
fibrous, sheetlike, and helical. Many hundreds of
these proteins are known in the insect group alone.
In some aquatic arthropods, such as crabs and
other crustaceans, the chitin and protein are
accompanied by minerals, especially
chalky crystals of calcium carbonate.
These make the exoskeleton or
shell harder for protection. It also
makes it heavierwhich is of
less hinderance to aquatic
animals whose weight is
supported by water
and consequently also
more brittle and liable
to fracture.

sticky feet
A jumping spider has two
foot claws and a pad of
tufted hairs that are formed
by tiny, supple extensions of
the cuticle. These can stick
to a wide variety of surfaces.

new armor
A spider crabs heavily
mineralized shell is hard
and rigid, except for
where it thins at the
joints. During the few
hours after each molt,
the soft new cuticle
enlarges rapidly before it
hardens. At this time the
crab hides away because
of its increased vulnerability
until its new shell is ready.


many segments make

up caterpillars body

a wasp has a typical insect

exoskeletonwith a head,
thorax, and abdomen.

wood lice are terrestrial crustaceans

with a shieldlike segmented
exoskeleton for protection.

spiny bristles deter


a caterpillars thinned body

cuticle allows each segment
to flex and change shape.
forewings modified
as rear-wing covers

a rhinoceros beetles
horn is formed from
thickened, stiff cuticle.

Structure and layering

A typical cuticle has several main layers. The outermost layer, the epicuticle,
is the barrier to the outside worldrepelling microbes, coping with physical
wear, and reducing water loss. The procuticle is formed from chitin fibers
and mineral crystals embedded in a variable matrix of proteins. Neither of
these layers contains living cells. The epidermis beneath these is the layer
of living cells that manufactures the top two layers. Under the epidermis,
the basement membrane forms a firm support with fibers of the protein
collagen. The relative
proportions, compositions,
and strengths of these
layers vary between
arthropod species, and
also on different body parts.
The procuticle layer is divided into the
hardened exocuticle, which has many
compacted fibers, and the more flexible
endocuticle. Dermal glands in the
epidermis can produce chemical
repellents to deter predators.

gland cell


Chalky skeletons
The bodies of some invertebrates are supported by chalky frameworks.
These can take two forms. Mollusks, such as snails, tend to have shell-like
coverings, while echinoderms (including starsh and sea urchins) are
more likely to have what can be termed a true skeleton. In both groups,
a distinctive feature of their structure is calcium carbonate, the main
constituent of rocks such as chalk and limestone.

Chalky body structures are usually laid down in the form of calcium crystals
embedded in a matrix, which is usually protein based. Calcium carbonate
may be joined by allied minerals such as calcium phosphate, magnesium
carbonate, and silicates. Also, calcium carbonate itself crystallizes in a variety
of forms, such as angled, prismlike calcite and more rectangular aragonite.
Marine mollusks tend to have calcite crystals, while argonite is more common
in terrestrial mollusks, such as land snails. The predominant material in
echinoderm skeletons is calcite, whose crystals tend to lie in the same
orientation within each of the small skeletal elements, called ossicles. These
are almost bony in texture, being hard and stiff, and spongy rather than solid.

Echinoderm means spiny skin, as
exemplified by the crown-of-thorns
starfish, shown here feeding at
night on coral in the Red Sea.

This triton trumpet is seen feeding on a crown-ofthorns starfish. The shell of this sea snail is made
of calcium carbonate, which it gets from its diet
and the surrounding seawater.

The flexible arms of the snake starfish,
a type of brittlestar, writhe like worms
because their ossicles are loosely bound
into the pliable body wall.



ball-andsocket joint

ossicle plate

The standard echinoderm arrangement is a thick outer

body layer containing many embedded calcium-based
ossicles. These vary greatly in shape and sizefrom
microscopic to palm-sized, depending on not only the
species but also on the body part concerned. This
type of skeletal structure is technically dened as an
endoskeleton since it is not produced by the outermost
body layers. But it functions as an exoskeleton because
it encloses the main body parts. In urchins the skeletal
plates are large and locked together to form a rigid
ossicle plate
covering. In more exible types
of echinoderms, such as sea
cucumbers and brittlestars,
The inflexible ball-shaped skeleton of a sea
urchin is known as a test. Joints attach the
the ossicles are embedded
spines to the test. The waving tube feet,
in a matrix of proteins and
part of the water vascular system, protrude
through small holes in the plates.
other substances.

tube feet (cover
whole body)


This microscope image shows a collection of sea
cucumber ossicles (shaped like sheets, wheels,
spines, buckles, and crosses) and silica-based
slivers (called spicules) from a sponges skeleton.



The highly magnified
ostracum of an abalone
shell reveals layers of
overlapping, platelike
aragonite crystals.

A mollusk shell is secreted by a gland in the mantle, which

is the animals eshy, cloaklike body covering. Shell form
varies hugely, with segmentlike plates in chitons; two-part
shells in bivalves, such as mussels and clams; a winding
helix in gastropods; and a much reduced internal structure
in cephalopods, such as squid, or no shell at all in
octopuses. The shell includes a thick central layer, the
ostracum, itself composed of two
layers of calcium carbonate crystals.
The nautilus adds
inner walls, or septa,
On the outside of the ostracum is the
to its shell, and lives
protein-rich periostracum that protects
in the last, largest
the calcium carbonate layers of the
chamber. Chitons, or
coat-of-mail shells, have
ostracum from dissolving or chemical
eight sectional plates
attack. The innermost shell layer
and a surrounding
is the aragonite-rich hypostracum,
girdle skirt. Murex
shells are among the
which in some mollusks has a lustrous
most complex of
sheen and is commonly known as
gastropods and of
all mollusk shapes.
mother-of-pearl, or nacre.
coiled shell











A snail shell gradually grows in diameter
as new material is added, and its size
reflects the snails age and food supply.


A bivalve shell grows as the soft, fleshy
mantle adds new carbonates and other
substances around the edge.

A squids shell is wholly internal
and known as a pen. It is thin,
lightweight, and pliable.

mantle cavity

A squids internal shell serves as an endoskeleton,
helping the body stay relatively stiff and giving
the muscles of its water-jet propulsion system
a firm base for their contractions.


Bony skeletons
Most vertebrate animals have an internal framework (or endoskeleton)
consisting chiey of the hard, mineralized tissue known as bone. It is
divided into the axial skeleton and the appendicular skeleton. The axial
skeleton runs along the middle axis of the body, from head to tail. The
appendicular consists of the bones attached to the axial skeleton.

Sharks, skates, and rays have skeletons, but
these are made mainly of cartilage rather than
heavier bone. As a result, swimming is more
thoracic vertebrae

cervical vertebrae







lower canine

Bone and cartilage


Under a bones outer hard
layer, cancellous bone tissue
(above) has many tiny
chambers. Cartilage (here
colored green) is light and
pliant, forming support for
body parts such as the ears.

Bone is a complex tissue, composed of mineral crystals (mainly

calcium phosphates) along with brous proteins, principally
collagen, embedded in a ground tissue or matrix of carbohydrates,
salts, and other substances. It is continually maintained by cells
known as osteocytes, which can also repair damage, such as
fractures. A typical bone has a dense, hard outer layer of
compact bone tissue; a layer of cancellous bone tissue beneath
this, which is more spongy or honeycomb-like; and a central
cavity of jellylike marrow, which stores fat and manufactures new
blood cells. Cartilage is similar to bone, with protein bers,
carbohydrates, and other substances encased in a matrix.
However, it lacks the hard calcifying minerals of bone, making it
somewhat lighter, softer, more pliable, and less brittle.

Skeletal appendages
In addition to an internal skeleton, some
vertebrates have associated bony parts and
appendages that develop in the same way as
the skeleton, but elsewhere in the body. Some
shes, such as armored catshes, have stiff,
bony plates in their skin. Their decreased
mobility and increased weight are offset by
greater physical protection. Reptiles such as
the draco, or ying dragon, have long, thin
rods of bone that hold out aps of skin.
Among mammals, physical protection in
armadillos is achieved by hard bony plates.
These form crosswise bands around the
body and consist of dermal bone, which
has no links with the endoskeleton. It forms
within the thickness of the skin and is
covered by a layer of horn-coated, bonebased scales, known as scutes.

Each of these animals possess
bony parts or appendages that
have developed for varying
functions, such as protection and
self-defense. The sea horse has
developed bony plates in its skin for
protection against predators. The
neck frill of the frilled lizard is spread
out when the lizard feels threatened,
and the bony plates and scutes of an
armadillo can protect its whole body.




Sea horses have an outer layer of bony

plates covered by thin skin.


The frilled lizard spreads out its wide

neck frill to deter enemies.

Three-banded armadillos roll into

balls for all-over body protection.


lumbar vertebrae

sacral vertebrae


The 200-plus bones of the tiger skeleton form a
lithe, flexible inner framework for more than 600
muscles to pull and maneuver. The vertebral
column runs from the skull to the tail tip. It acts as
a flexible central support for the limbs and ribcage.
In mammals such as cats and dogs, the skeleton
forms about 20 percent of the bodys total weight.

Each bone of the tigers skeleton
has a part to play in its movements
and behavior. Opening the jaws to
snarl, swishing the tail, and crouching
ready to spring are all achieved by
teams of muscles pulling precisely
on particular bones.



costal cartilage

Highly modied skeletons


The typical vertebrate skeleton consists of a skull,

a exible backbone, and four movable limbs, but
through evolution the basic layout has become
hugely modied for different habitats and lifestyles.
In snakes, only the skull, greatly lengthened vertebral
column, and ribs remain, while most or all limb
remnants are lost. Turtles and tortoises have a rigid
enclosing shell made of the domed carapace over the
back and the atter plastron on the underside. In birds,
the forelimbs are greatly altered as wings for ying, with
reduced numbers and sizes of bones, especially the ngers.
Whales and dolphins have forelimbs modied as ippers,
but lack any rear limb bones. In bony shes, bony rods,
called rays, hold out the exible ns.


caudal vertebrae



All snakes have more than 100

vertebrae, and some possess over 400.

Swimming muscles anchor to the long

extensions of a shs vertebrae.

Each of these vertebrates has a bony skeleton with
a skull, spinal (vertebral) column, and ribs. But the
other parts and proportions of the skeleton vary
hugely, for functions such as flight, swimming,
slithering, and self-defense.

The turtles ribs and vertebrae are

fused to the inside of the carapace.

Bird bones are mostly thin, hollow,

and light to allow for ight.




Cheetahs require fast, brief bursts of speed to
catch their prey. They have an extremely flexible
spine, which coils and uncoils with every
movement, propelling the animal forward. The
cheetah can reach speeds of 70 mph (113 kph),
but they are not built for stamina. They become
breathless and overheated within 30 seconds.


Animals display an incredible variety of movements, from the hovering
of hummingbirds to the slithering of snakes. Locomotion occurs when an
animals entire body moves from one place to another, as when walking,
running, swimming, ying, leaping, or crawling. In addition to utilizing
muscle power, animals can also harness a number of environmental
features, such as the wind, water currents, and gravity, to aid movement.
The desert arachnid called the solifuge (also known
as the sun spider) has a clever method of escaping
predators. It curls its legs around to form a wheel
shape and rolls down sand dunes.

Land, air, water, and soil


Methods of locomotion vary dramatically, according to the substances

or media an animal is traveling through. Moving across the land
necessitates contact with the ground, using body parts that range from
legs and feet, to scales in snakes, and a slimy undersurface or foot
in snails. Grip against the surface provides the required forward thrust.
The huge variety of surface consistencies requires numerous
specializations. For example, desert dwellers such as kangaroo rats
have enormous feet and toes as well as hairy soles to help them jump
in the soft, shifting sand. Moving through air requires large aerodynamic
surfaces, such as wings in true iers (birds, bats, and insects), or aps

Various animals take advantage of

gravity, by tumbling or sliding down
slopes, or simply falling through the air. These methods are generally
used as emergency measures to escape predators. A number of animals
possess the ability to curl into a ball and roll away from danger, including
millipedes and wood lice. Tree-living insects, such as beetles and stick
insects, simply let go and drop to the ground. Their survival is ensured
by their small size, tough body casing, and soft landing site (leaf litter).

Metabolic rate
There is a close connection between metabolic rate (the speed of essential
biochemical processes within an animals body) and locomotive ability.
Birds and mammals are homeothermic (warm-blooded)that is, they
maintain a constant high body temperature. This makes their muscles
ready for action at any time. Most other animals are ectothermic (coldblooded), so their temperatures vary according to the environment. When
the temperature cools, their muscle metabolism becomes less effective.
At very low temperatures, they are actually unable to move at all.






of skin in gliders, such as colugos and ying
Swallows have scythelike wings
squirrels. Animals have to expend a lot of
for speedy, acrobatic pursuit of
energy to provide both lift (to counteract
food. Earthworms are long and
slim, to push between soil particles.
gravity) and thrust for forward movement.
The jellyfishs shape exploits sea
Water is a much more resistant medium
currents, and the kangaroos hops
than air. In general, it requires at least twice
are energy-efficient on soft ground.
the amount of energy to attain the same
speed in water as it does on land. A smoothly contoured, streamlined
shape becomes extremely important in water. Aquatic creatures generally
have large, at surfaces such as tails and ns to push against the heavy,
uid medium. Locomotion in soil, sand, and mud is by far the slowest
and most energy-intensive. However, as in all forms of locomotion, there
are benets to offset even the
most arduous movement. For
example, burrowing animals are
usually less visible to predators
and are sheltered from the
weather. They may also be
surrounded by their common
food source, such as plant roots
in the case of naked mole rats.
Some appendages are a compromise between
various forms of locomotion. The mudskippers
muscular pectoral fins can help this fish
burrow, swim, wiggle, waddle, and leap.


On cool evenings, some hawkmoths have to
shiver their flight muscles to generate enough
heat to take off.

The swordfish is able to channel natural low-level
heat within its body along certain blood vessels to
warm its muscles, enabling short bursts of speed.

This chart shows four mammals of similar massesabout 134 oz (50 g) that move through
land (eld mouse), air (bat), water (desman), or soil (mole). It plots how far each travels in one
second and how much energy it expends (for comparison, the eld mouses energy expenditure
is shown as one unit). The bat uses energy most efciently, and the mole the least.
Distance traveled in 1 second (meters)

Units of energy used per second compared to field mouse














anImal anaTomy 90

Walking and running

(heel bone)

With the exception of a few extraordinary animals that are able to walk on water or
along the ocean floor, walking and running are confined to land animals and require
the use of legs. There are many different limb actions that animals employ to achieve
locomotion on land, which are known as gaits. Individual animals often exhibit a
broad variety of gaits. These methods of locomotion are most obvious in larger
mammals, but they also occur in the smallest invertebrates.

(heel bone)


Different gaits

The numbers of legs varies greatly through the animal kingdom, from
two in birds and kangaroos, to the standard vertebrate number of four
in most mammals, amphibians, and reptiles. Insects have six legs, while
arachnids and some crustaceans have eight. Centipedes often possess
well over 100 legs. Despite their name, millipedes do not ever have a
thousand legs. Most species have between 100400 legs, but onethe
Illacme plenipeshas an incredible 750. In centipedes and millipedes, the
sheer number of legs is an arthropod adaptation to a part-burrowing way
of life. The many tiny limbs allow these creatures to push powerfully
through loose material, leaf debris, and soil, without the individual limbs
needing a lot of space for their actions. The legs move mainly to and fro
rather than out to the side, which keeps their movement efficient.

Many mammals use different gaits,

from a slow walk to a full-speed
run. These gaits are especially
clear in hoofed mammals, such
as horses, which alter their leg
movement from walk to trot,
canter, and gallop. The trot is used
to cover long distances efficiently,
while the gallop is used to evade
predators. Unusual gaits include
camel pacing, where both legs on
one side move simultaneously, to
give a side-to-side swaying motion.

fatty pad
foot bones


In many creatures, specialized body

movements aid locomotion. For
example, the flexible backbone of the
cheetah arches up and down to extend its
stride. In salamanders, newts, and lizards
(among others), S-shaped sideways curves
pass along the body from head to tail. This
gives added swinging motion to the limbs,
which splay out to the sides.

Leg design

A tiger salamanders body
curls from side to side as
it walks, in a series of
S-shaped waves.
This pattern is
derived from
the movements
of fishy ancestors
of amphibians.

compromise limBs

Rhinoceroses have thick, sturdy limbs to carry their great bulk.

In spite of this, they are able to sprint surprisingly quickly.

Rhinoceroses and ostriches
rely on their muscle mass
close to their main body to
achieve locomotion. Starfish
locomote by hydraulic
pressure. They squeeze
water into each of their tinytubed feet to extend them.

Ostriches contain their main musculature in their hips and thighs.

They can also use their wings as rudders to help change direction.



Plantigrades, such as bears, walk with their heel bone,
metapodials, and digits on the ground. Digitigrades,
such as dogs, move with only their digits touching the
ground. Unguligrades stand on one or more toe tips.
Externally, elephants appear plantigrade, but are
actually digitigrades, since the heel bone is raised and
only their digits touch the ground.

Specialized movements

In general, longer limbs allow longer strides

and greater speed. With fast runners, the
muscle bulk that moves the limb is near the
main body, often in the shoulder or hip region.
This reduces the weight of the limb toward its end,
making it easier to move to and fro at speed. Many
invertebrates have unjointed legs, which often utilize
hydraulic pressure. Caterpillars have both jointed and
unjointed legs. The latter help the caterpillar to grip,
since they contain tiny hooks that act like suction cups.



Numbers of legs

A crabs sideways walk is due to the
direction in which its leg joints bend. A
millipedes legs move in coordinated waves
a single wave involves lifting around a dozen legs
up, lowering them, and then pushing backward.


Starfish have tiny tubes on the underside of their five

or seven arms, which lift up and move forward.

Animals that move both on land and in

water have compromise limb designs.
Inter-toe webbing is an enhanced version of
the standard five-toed land vertebrate foot.
It provides a broad, finlike surface for
pushing against water and is found in
a wide variety of semiaquatic walkerswimmers, including otters and desmans
among mammals, many kinds
of seabirds and waterfowl
from albatrosses to
ducks, and many
The degree of
webbing reflects
the proportion of
time spent in water
for example, tree frogs
have virtually none.

Ostriches can sustain a speed

of 45 mph (70 kph) for up to
30 minutes, covering up to
161 2 ft (5 m) in a single stride.

As a zebra gallops, all four hooves are off the
ground for more than half of the time taken for each
complete stride. Such minimal contact reduces
friction with the ground and allows the zebra to
fly in a succession of long leaps, at speeds
exceeding 35 kph (55 kph).


Climbing and leaping

There is a huge diversity of climbing animals, some of which have developed
extraordinary specializations, such as the acrobatic skills of gibbons or the ability of
geckos to stick to almost any surface. Leaping involves progression by alternately
speeding and slowing, in a series of jerky actions. Most animals use the same limbs
for leaping as they do for walking and running, although some invertebrates use
additional body parts specifically to leap.

Getting a grip
Moving through tree branches, and up cliffs, rocks, and walls requires
strong, mobile limbs and an excellent grip. Most climbers have muscular
limbs that can haul their body weight upward. Since it is necessary to
hang on while the other limbs are moved to new positions, some climbing
animals can support their body weight with only two or even one limb
gripping. Powerful claws, fingers, toes, and tails
contain various specializations to achieve grip. For
example, a chameleons five toes are grouped as two
sets, of three and two, to form a pincer that clings
onto a twig with a vicelike hold. Some animals,
including a number of monkeys, possess prehensile
tails, which are able to grasp and hold objects.


Geckos have ridged toes (left) with thousands of
minuscule stalklike bristles, dividing into billions of
microscopic spoonlike hairs (right). These hairs
mesh with the tiny irregularities of the surface the
gecko is climbing and provides it with grip.

The tip of a tarantulas foot
has two claws, a hook,
and serrated hairs, all
of which grip strongly
to most surfaces.



Primates have adapted limbs not just for
locomotionchiefly in treesbut also for
feeding and grooming. The chimpanzees
muscular, semi-opposable toe and thumb grip


branches very well. The greatly elongated

middle fingers of the aye-aye pick out grubs
from under bark. The indri lemur has evolved
a grooming claw on its second toe.




A proboscis monkeys feet are

partially webbed, making them
very effective swimmers.


Locomotion by brachiation (arm-over-arm swinging)
occurs chiefly in tree-living primate mammals,
and most spectacularly in gibbons. There
are 14 different species of gibbons, which
are found in the tropical rain forests of
Southeast Asia. The siamang (right) is the
largest of them. It is perfectly equipped for
brachiation, having long-palmed, hooklike hands
with much-reduced thumbs, elongated, powerful
arms, muscular shoulders, and flexible joints.
This is aided by stereoscopic distance-judging
vision, which helps locate the next handhold.
When swinging, the siamangs body moves forward
in a series of arcs, like a traveling pendulum. It is an
energy-efficient form of locomotion because it involves
maintaining momentum.






1 At the start of each swing, the siamangs body
gains speed and energy as it swings forward.
2 Its body swings around to allow the free hand to
grab the next branch. 3 The long, muscular arms
and the momentum from the swing propel the
siamang to the next branch. 4 The grasping feet
reach for a lower branch as it comes to a halt.

Energy-efficient locomotion

Some species of monkey, including
the proboscis monkey, make their
longest leaps to escape danger.
Both fingers and toes are adapted
for grasping, and the tail functions
as a balancing rudder.

How animals leap

Prodigious leapers, including, hares, kangaroos, frogs, fleas, and
grasshoppers, usually have one pair of specially adapted limbs that are
larger and stronger than the others. Each limb unfolds sequentially at its
joints as a series of levers, from the hip and thigh, to the knee and shin,
to the ankle and foot, and lastly, to the toes. This flings the animal up
and forward in a series of leveraged pushes, achieving a rapid gain in
momentum. In addition to avoiding predators and other dangers, leaps
are used in many other contextsfor example, to clear obstacles, reach
a nearby branch, or as a display of fitness when courting or defending
territory. Desert-dwelling animals often use short, quick leaps as an
efficient way to move over soft, shifting sand.
When on the ground,
some lemurs and sifakas,
such as this Verreauxs
sifaka, move with
sideways bounds of their
long, muscular legs. They
raise their arms up and
outward for balance. This
remarkable form of
locomotion can propel
the sifaka over 16 1 2 ft
(5 m) in a single leap.

The energy efficiency of leaping is improved by

structures in the limbs, such as ligaments around joints,
and tendons, which anchor muscles to the skeleton.
These structures contain rubbery,
elastic substances, including the
proteins elastin and resilin. As
ligaments and tendons stretch or
compress in preparation for the
leap, they store energy like a coiled
spring. The energy is then released
during the leap, in a catapult action
that assists the muscles. As a
kangaroo lands after even a small
Before a jump, this cat
hop, its body weight stretches its
flea compresses blocks
leg tendons and ligaments, storing
of resilin at the bases
of its legs.
energy for the next hop.

13 in

The leap of a 1 16 in
(2 mm) cat flea170 times
its own body length.
Certain hoofed mammals perform stiff-limbed, springlike vertical
leaps, as though bouncing along with their legs held straight. This
motion is known as pronking and is derived from the Afrikaans word
pronk, meaning to boast. It occurs particularly in antelopes, such
as the springbok (shown here) and impala, as well as many gazelles.
This behavior is thought to be a
display of fitness, showing
predators that the
individual is healthy,
and not worth


Burrowing, slithering, and sliding

Some forms of locomotion involve progress in tiny, often
continuous stages, with a large area of body in contact
with the surface. These movements include sliding
and slithering, which are undertaken mainly by limbless
creatures. Many animals are capable of burrowing and
this occurs through many different types of terrain.

Burrowing and tunneling

There are many subterranean animal species that
employ a variety of methods and body parts to push
aside particles of soil, mud, sand, or similar material
and force themselves forward. This is by far the slowest
and most energy-expensive method of animal
locomotion. However, there are benefits of the fossorial
(underground) mode of life for creatures that habitually
spend their lives tunneling or burrowing. A truly
fossorial animal is relatively safe, since it is out of sight,
hearing, and scent of surface predators. It is also
sheltered from extreme conditions, such as droughts
and blizzards. Also, many burrowing creatures exploit
underground food sources such as roots, bulbs, and
other subterranean plant parts.

65 ft

The length of tunnel

an average European mole
excavates in a single day.
The sharp claws on the European
moles enormous front feet work like
shovels. The mole anchors itself with
its back feet and scoops soil sideways
and backward. When it nears the
surface, this shoveling action pushes
the soil up, forming a molehill.

Piddocks are bivalve mollusks that have shells with

BURROWING ridged teeth to rasp into rock.

Some animals use
specialized body
parts to burrow through
hard surfaces, such as
wood and rock. Softer
substances also
present challenges.
Sand collapses as the
burrower passes, leaving
no permanent tunnel.
This means that creatures
like sandfish must Sandfish (a species of skink) wiggle like fishes to
continually expend swim through loose sand and soil.
energy as they go.

Shipworms are bivalve mollusks that possess tiny,

serrated shells, which enable them to bore into wood.

Naked mole rats use their large, constantly growing

incisor teeth to bite through dry soil.


Burrowing methods
A typical burrower needs to push aside particles of the
surrounding ground, using muscle-powered pressure
from one part of its body, while at the same time
anchoring another part of its body to generate sufcient
burrowing force. Vertebrates, such as moles, have
powerful limbs that work like shovels to shift soil. Many
invertebrates, such as subterranean termites, have sawtoothed mouthparts, which they use to cut through the
soil. Some bivalve mollusks, such as the razor shell,
have muscular feet, which expand and contract to
enable it to burrow through mud and sand. Aided by its
streamlined shape, which minimizes resistance, this
burrowing method allows the razor shell
to dig 3 ft 3 in (1m) in under 10 seconds.

Slithering and sliding

A slug travels using small waves of muscle
contraction, passing from head to tail along its
broad, slime-coated foot, pushing it forward
like a conveyor belt.

Snakes, slugs, snails, atworms, and similar animals

move smoothly and continuously along a surface, as
a result of many tiny muscular contractions. Some
snakes tilt their scales to gain purchase against objects and
undulations on the surface. Gastropod mollusks, such as
snails, slugs, and limpets, slide on a lm of mucus using
rhythmic actions of their foot muscle layers. These actions
move the foot along in undulating waves. The suction achieved
by the sticky mucus allows them to grip
onto many different surfaces, including
If this garden snail moved
rocks and loose soil. It even enables
continuously, it would take it over
a week to cover 0.6 miles (1 km).
them to travel upside down.


The razor shell pushes its valves
(shell halves) slightly open and
extends its long, fleshy foot into
the sand to thrust downward.

The foot swells at the tip to form a
lower anchor point, then the foot
contracts, the valves close together,
and the shell slides down.

In contrast to animals with legs, snakes have no

concentrated point to push off from. Instead, they have
a complex system of muscles, which allows them to move
using four distinct methods. Often, the method used varies
according to the size of the snake, the kind of surface they
are traveling on, and how quickly they need to move. Most
snakes can perform each of these types of locomotion as
the situation arises. Sidewinding is the only exception,
which is unique to the caenophidian family of snakes.

In rectilinear locomotion, the belly scales are lifted, tilted to grip the
surface, then pulled forward, in a succession of waves along the body.
This method is mainly used by heavy snakes.

In concertina locomotion,
the rear body folds in sideways
curves, which act like a
frictional anchor. The head
then extends forward, and
the rear is drawn along.

Most species of snakes,
including this Burmese
python, are able to vary
their style of locomotion
according to the terrain
they are traveling on.
This python is utilizing
concertina locomotion,
most often used when
climbing, or traveling
through tunnels.

In sidewinding,
the snake lifts its
head and extends
it forward. The
rear of the
snakes body lies
side-on to the direction
of movement, allowing for
better leverage. This is
employed on smooth or
slippery surfaces.

In lateral undulation, the snake

exploits surface irregularities, such
as rocks, tree trunks, and hillocks,
by adjusting the angle of contact
to gain forward thrust.


Flying and gliding

The three main animal groups to have mastered the air are insects, birds, and bats.
They are considered to be true iers, since they all capable of staying airborne by
apping their wings. Some other animals are able to employ temporary airborne
locomotion but they are considered gliders because they are not capable of powered
apping ight. Gliding is also employed by some of the true iers as an energy-efcient
method of locomotion. Some birds are able to use thermals (columns of warm air) to
soar for hundreds of miles. This means that they hardly ever need to ap their wings.

Insect ight

Bird ight

Almost all the main insect groups have the power

of ight. Typically they are four-winged, such as
dragonies, mayies, butteries, caddis ies, moths,
bees, and wasps. In the true iesmore than 120,000
species of houseies, blowies, gnats, midges,
mosquitoes, hoveries, crane ies, and othersthe
hindwings have actually become tiny, drumsticklike
secondary wings. These vibrate or twirl rapidly, adding
stability and control, while the forewings actually
provide lift and thrust. The ight muscles are located in
the thorax (middle body section). Some insects directly
contract and relax these muscles to pull the wing bases
up and down. Other insects actually change the shape
of their thorax in order to move their wings up and
down. For both techniques, muscles at the wing bases
determine the direction of ight by adjusting the angle
of each stroke. Wing motion is not just up or down,
but also to and fro, to generate both lift and thrust.

Bird wings create much of their lifting force by forward

movement through the air, using the airfoil design (see
panel, right). The main power for apping comes from
the pectoralis major (breast) muscles in the chest.
These anchor to the sternum (breastbone) at the center
of the chest, and at the other end to the inner wing
bones. As these muscles contract, they pull the whole
wing down and back. Muscles within the wing, aided
by long tendons running out through the leading edge,
can ex or warp the whole wing, change the angle
of the feathers, and alter the wings curvature for
precise control.

A birds wing forms a curved shape along its
upper surface, known as an airfoil. Air
passing over the upper surface has to travel
slightly farther at a faster pace than the air
passing along the lower surface. The slowmoving air beneath the wing exerts a
greater pressure, which effectively pushes
up the wing from below. The faster-moving
air above the upper surface of the wing
produces lower air pressure and sucks the
wing up from above. This creates a
continuing force of lift that true iers need
to counteract the downward pull of gravity.
low air

faster air

slower air

high air

cross section
of birds wing


A barn owl raises its wings on the upstroke.
It twists its feathers to allow air through, thereby
reducing resistance. It then lowers its wings
strongly on the powerful downstroke. The feathers
flatten to overlap and form a continuous airproof
surface. This ensures maximum lift and thrust.

Among the fastest,
most aerobatic insects
are dragonflies. They
can rapidly accelerate
to speeds of more than
40 mph (65 kph).
Dragonflies are also
able to beat their two
sets of wings alternately,
which increases


flexor muscles

biceps muscle


primary flight

In beetles, such as
ladybugs, the forewings
have become hard, protective
covers, called elytra. Before
takeoff, these are raised and held
clear as the larger, delicate, hind
wings unfold for flight.


The primary feathers fan or fold to control
speed and direction. The secondary
feathers form the main airfoil for lift. The
alula (digit covered by 35 feathers)
disrupts smooth airflow at the leading
edge to reduce speed for landing.

secondary flight




Gliding specialists
There are a number of animalssuch as ying squirrels, ying possums,
ying lizards, ying frogs, and ying sheswhose names suggest they
can y. They are not, in fact, true iers, since they cannot remain airborne
in a sustained way and gain height under their own power. These
remarkable creatures employ winglike structures with large surface areas,
which function as parachutes to increase air resistance. They can also
generate small amounts of lift that reduce descent speed. Tilting or
changing the shapes of these surfaces gives a certain amount of control
over distance and direction. The most impressive mammal gliders are the
ying lemurs (neither true iers nor true lemurs) of Southeast Asia, which
can travel more than 330 ft (100 m) and land with pinpoint accuracy.
When the sun warms the
ground, the air above it
also warms. This creates
columns of warm air,
known as thermals.
Updrafts are formed in
mountainous terrain
by deflected wind. Both
thermals and updrafts
enable birds to soar
with minimal need to
flap their wings. This
energy-efficient method
of locomotion is used
by eagles, vultures,
condors, and storks
among others.



California flying fish use their pectoral and pelvic
fins as gliding surfaces. Wallaces flying frog
extends its webbed toes as four mini-parachutes
to slow its fall, as it jumps from a tree.

Wing aspect ratio

A wings aspect ratio refers to the relative proportion of its span
(length from the body to the wing tip) to its width (from leading to
trailing edge). Long, narrow wings have a high aspect ratio and
are most effective for long-distance gliding and soaring. Flapping
is minimized, since the air currents provide much of the lift
needed to sustain ight. Many birds feature this wing design,
including albatrosses, gulls, and to a lesser extent condors and
eagles. Low aspect-ratio wings are shorter and wider. These are
used for rapid acceleration, fast
turns, and precise control, as in
sparrowhawks. Curved,
scythelike wings with
tapering, drag-reducing
tips are used for sustained,
speedy aerobatics, as seen
in swallows and swifts.
Hummingbirds flap their wings in a figure-eight
pattern, providing the necessary downflow of air to
allow them to hover. With slight adjustments, they
can fly sideways and even backward.




Swimming is remarkably similar to ying in a number of respects.
Both air and water are uid media, and many of the same principles
apply to ns as to wings, such as the need to push broad surfaces
backward in order to propel an animal forward. One signicant
difference is that water is 1,000 times denser than air, bringing many
drawbacks, but also benets.

Staying aoat
An advantage of water over air is that it provides plenty
of support, so unlike aerial animals, aquatic species do
not need to generate powerful lift. However, they do
need buoyancy control, as well as propulsion, in order
to rise, descend, or hover in the water at a certain
depth. Bony shes adjust their buoyancy using an
organ called the swim bladder, the gas content of
which can be adjusted. Cartilaginous shes, such as
sharks and rays, lack a swim bladder. Their angled ns
provide them with hydrodynamic lift as they move

Juvenile blackfin barracudas
often swim in large schools.
They have long, streamlined
bodies with very powerful
tails, which allow them to
accelerate rapidly.

forward, offsetting a tendency to sink. Sharks also

have a large, oil-rich liver that adds to their buoyancy,
since oil is lighter than water. In cephalopods, such as
nautiluses, cuttlesh, and squid, the shell (which is
internal in the latter two) contains gas-lled spaces that
can be lled or emptied to adjust buoyancy. Diving and
swimming birds have adaptations that enable them to
sink quickly in order to catch food. Cormorants, for
example, have feathers that can hold a lot of
water, reducing the air trapped in their plumage.

swim bladder


gas gland

rete mirable (network
of capillaries)

Each chamber of a nautiluss shell is filled with fluid, which is
absorbed and replaced by gas. An opening in each chamber allows
the nautilus to control the volume of gas inside, and therefore
regulate its buoyancy. The siphuncle removes excess water.

Swimming styles
Most large aquatic animals, such as marine mammals
and big shes, swim using the muscles in their bodies
and tails to push against the water. Some cartilaginous
shes, bony shes such as eels, and sea snakes swim
with an undulating or wriggling motion, as S-shaped
waves travel along the entire body. Most bony shes
swim by moving their rear body and tail (caudal n)
from side to side. The other nsdorsal on the back,
anal on the underside, the paired pectorals at the front,
and pelvics toward the rearare generally used for


A bony fish regulates its buoyancy by exchanging gas between its
bloodstream and its swim bladder. Gas is secreted into the swim
bladder from the gas gland, which is supplied by the rete mirabile.
The gas is removed and reabsorbed into the blood by the ovale.

steering and to provide stability. However, some

shes use them for their main propulsion, including
seahorses, which swim by rippling the dorsal n, and
rays, which undulate their large, winglike pectoral ns.
A shs body shape usually reects its swimming style.
Powerful, fast movers have muscular, torpedo-shaped
bodies that are elongated and tapered at both ends.
Fishes that live in quiet waters and do not need to
move at high speeds, except in short bursts, tend to
be laterally compressed (narrow from side to side).
Bottom dwellers are usually vertically compressed,
(narrow from top to bottom) for example atshes
and angel sharks.

A sharks tail n has an

uneven shape, generating
both lift and thrust.

Seals like the Hawaiian monk seal move by
undulating the rear body and kicking with
their rear flippers. Eels, such as the whitemargined moray, utilize serpentine locomotion
moving in a series of muscular waves
passing from head to tail. This is aided by a
flattened tail, which generates more thrust.

A tunas tail n lowers

drag for fast cruising.

Bony shes have a

symmetrical homocercal
tail n.

The shape of a
fishs tail reveals the
swimming style of its
owner. Most bony
fishes have a
homocercal tail
shape, with upper
and lower tail lobes
approximately equal.

A pikes tail n has a large

surface area for fast acceleration.

of fish

of tail

resultant force

As a fishs tail moves, it generates both sideways
and backward thrust in the water. The resultant
force acts diagonally halfway between the two.
As the fish moves its tail from side to side, the
resultant diagonal thrusts to left and right produce
a net backward thrust, so the fish will swim in
a straight line.

Some smaller animals propel themselves
through water using tiny hairlike structures.
Comb jellies have eight rows of tiny hairlike
cilia that beat in coordinated action, like
miniature oars. Some diving beetles (below)
swim with their hind legs working
simultaneously to push water backward.
Their legs are fringed with hairs, which
open out to effectively form two paddles.

Legs and limbs

Jet propulsion

Many vertebrates other than shes swim using their

limbs. Seals propel themselves by wiggling the body
and clapping their broad, well-webbed hind ippers
together, while sea lions use mainly their front ippers.
Birds swim by apping or rowing with their wings or
kicking with their feet, or both. Penguins, however,
possess unique wings. Their bones are solid instead
of hollow, increasing their density and strength. They
do not row as with oars, but ap their wings and y
underwater. Marine turtles swim in a similar way,
but they use their front limbs
solely for propulsion and
Aquatic species like the kittiwake
their hind limbs to steer.
and caiman have webbing between
Some species of turtle can
their digits, which increases thrust
reach up to 18 mph (30 kph). due to the larger surface area.

A specialized form of aquatic locomotion known as jet propulsion is seen

in various cephalopod mollusks, including octopuses, squid, cuttlesh, and
nautiluses, and some species of jellysh. In a squid, the mantle muscles
relax, slowly drawing the water into a chamber called the mantle cavity
between the animals main body and its eshy,
cloaklike mantle. The mantle cavity expands
to accommodate the water. Muscle
contraction prevents water from escaping
the way that it entered. A powerful muscular
action then squirts the water out very quickly
through a funnel-like valve, called the siphon.
The cephalopod is pushed forwards as the
Cephalopods like this bigfin reef
squid use jet propulsion primarily to
water jets out.

Frogs swim by pushing against the water with
a synchronized motion of their hind limbs and
webbed feet. The whole process happens
quite slowly. The frog draws its legs up to its
body by bending the hips, knees, ankles, and
toes. Then is straightens its legs, with its
webbed toes spread, to ensure maximum
thrust. The human swimming style
breaststroke may have actually originated
as an imitation of the way a frog swims.


intake of water

allows water
to enter

escape danger. They can also move

slowly by rippling the fins along the
sides of their bodies.

mantle cavity

mantle cavity

muscle contraction
stops water escaping



The squid draws water into the mantle cavity using its
mantle muscles. The cavity expands to accommodate the
water. Sudden muscle contractions thrust the water out
through the siphon and propel the squid through the water.

expulsion of

The exquisite layout and patterning of the
differently sized scales around the eyes of a
reptile, in this case, a green iguana, allow
sufficient flexibility for the eyes to blink but still
provide a high degree of protection.


Animals and their environments are endlessly varied, and so are the
interfaces between themtheir body coverings. The outer surfaces, casings,
or skins of animals are highly adapted to their surroundings and vary from
microscopically thin and fragile to thicker and more durable than a brick.

Types and functions

An animals outer layers are known as its integument, or integumentary
system. These coverings range from basic skin and its outgrowthsscales,
feathers, hairs, and bristlesto horny sheaths and rock-hard shells. Each
covering is a complex compromise for a wide range of functions. Among
the most important are the containment and protection of soft inner tissues,
and the retention of body uids to maintain the internal environment. The
covering may also be involved in temperature regulation, nutrient uptake,
waste removal, gas exchange for respiration, touch and other sensations,
and features of appearance, such as camouage, warning colors, or mating
displays. These functions are mixed and matched for each animal.
Apart from hard-shelled animals, chiefly mollusks,
most body coverings have some degree of flexibility,
even if it is limited to small areas around joints, as in
insects and crustaceans. This flexibility allows the
animal to move around, feed, and avoid danger.
Hairy, furry, or bristly coverings occur in a wide
variety of animals. Such coverings are characteristic
of mammals but are also found in worms, certain
insects, such as caterpillars (larvae of butterflies
and moths) and bees, and some spiders.
Chimpanzees have the exible pinkish skin and
outgrowths of hair that are typical of mammals.

The shells of tortoises, turtles, and terrapins
provide excellent protection but are heavy, limit
movement, and restrict lifesyle.

protection but fulll few other functions. Some coverings are more
temporary, such as the outer casings of crustaceans and developing insects,
and bird feathers. These are shed or molted at intervals, taking any damage
with them, and are replaced by new, intact coverings. Very temporary
coverings, which are not part of the body itself but are bodily products,
include the bubbly froth or cuckoo spit secreted by froghopper bugs.

Temperature regulation and respiration

In some animal groups, the body covering can be adjusted to aid control
of body temperature. This is especially important in homeothermic, or
warm-blooded, animals, which maintain a relatively high body
temperature. In cold weather, birds
uff out their feathers and mammals
plump up their fur to trap more
insulating air and reduce heat loss.
A body covering may also function
in respiration by absorbing oxygen
and giving off carbon dioxide. This
function may supplement organs,
such as lungs or gills, that are
specialized for respiration, as in
A typical amphibians skin is moist and thin enough
amphibians. The body covering
to allow gases to pass through it. This is important in
may also be the sole means of
the Surinam toad, which stays underwater for long
periods and so cannot use its lungs for breathing.
respiration, as in some worms.
Walruses have a fatty blubber layer under
the skin to insulate against the cold; in hot
conditions, extra blood flows to the skin
surface to help lose excess body heat.

Birds, such as this whooper swan, are the only living

animals to have a covering of feathers.

The African bush viper snake, like most reptiles,

has a exible scaly covering over its skin.

The sea mouse, a polychaete worm, has bristles

known as chaetae projecting from its surface.

A feather stars arms are covered with exible

projections, called tube feet, that gather food particles.

A primary function of the body covering is defense. The external surface
is the rst point of impact for insults against a creature, which include
physical damage, destructive chemicals or toxins, harmful microbes,
parasites, and radiation. The covering may be extremely tough and
durable and able to resist these problems long-term, as in the shells
of mollusks and tortoises. These shells provide extra strong, extra rigid


Skinas distinct from most animal shells
or similar hard body coveringsgenerally
provides all, or almost all, of the outer surface
of an animals body, forming a continuous,
exible, protective body covering. Typically, skin
is made up of several layers and it may produce
appendages and outgrowths, such as hairs, scales,
horny or bony plates, or feathers.

Types of skin
In everyday use, the term skin usually refers to the exible outer
coverings of animals, especially vertebrates (shes, amphibians,
reptiles, birds, and mammals), rather than the rigid outer casing
of invertebrates such as crabs. In many animals, much of the
actual skin is under a layer produced by the skin itself, such as
scales in reptiles, feathers in birds, or fur in mammals. Apart from
the face, large areas of exposed skin are uncommon, occurring
only in a few shes, amphibians, and certain mammals such as
whales, hippopotamuses, walruses, naked mole-rats, and humans.

Caterpillars have a thin, exible outer cuticle

(an exoskeleton) that is made mainly of chitin.

The Egyptian vultures face is

covered in yellow skin.

Moray eel skin is smooth

and has no scales.

The skin of an animal is adapted
to its environment and lifestyle.
For example, aquatic animals tend
to have smooth skin to reduce
friction with the water. Terrestrial
animals tend to show greater
variation in skin texture, and often
have specialized areas of skin on
different parts of the body, such as
the featherless, waxy cere on the
heads of some birds.

Whale shark skin is

The re-bellied newt has red patches on
up to 4 in (10 cm) thick. its underside that may startle predators.

Whale skin is almost hairless, A gorillas facial skin can reveal

especially around the blowhole. complex expressions.

Hippopotamus skin produces

mucus to keep it moist on land.

Basic structure

hair shaft

The skin of all vertebrates has a similar basic microscopic structure, even
though it may be thinner than this page or thicker than this book. There are
three basic layers: the outermost epidermis, which is made up largely of
keratin (a tough, brous protein) and continually renews itself; the dermis
under this, which consists mainly of collagen and elastin bres and
contains sensory nerve endings, sweat glands, and blood
vessels; and an innermost subcutaneous layer. The
relative thickness and consistency of each layer varies
from species to species, giving each one its unique
skin features. For example, in cold-adapted birds
and mammals, such as penguins, whales, and
seals, the subcutaneous
layer is thickened with
Hagfishes have many mucussecreting cells in their skin. In just a
fatty deposits that form
few seconds, these cells produce so
insulating blubber to
much slimy mucus that, when mixed with
keep in body heat.
water, it would fill a bucket.

blood vessel
hair follicle
nerve ending
sweat gland

The skin of all mammals is fundamentally similar.
It has three main layers (epidermis, dermis, and the
subcutaneous layer) and contains many specialized
structures, such as blood vessels and nerve endings.

One of the most important roles for skin is defense

of the body and its delicate internal structures. This
defense may be physical, chemical, visual, or a
combination of the three. In mammals such as elephants
and rhinoceroses, the main physical defensive barrier is the
epidermis, which is hugely thickened and strengthened with
keratin. This tough protein occurs in various forms in many
vertebrates (as well as some invertebrates), and is also the
main component of hairs, claws, hooves, nails, horns, bills, and
feathers. Chemical defense is particularly common in amphibians;
in many species, the skin has tiny glands that ooze unpleasant or
poisonous secretions. Visual defense may take
the form of vivid coloration, which may prevent
The cane toads skin secretes a poisonous
predation, or camouage to help an animal
chemical. The wombat uses its leathery, thickconceal itself by blending into the surroundings. skinned rump to block its burrow against predators.

Nutrients and wastes

In some animals, especially soft-bodied invertebrates, the skin
acts as a selective two-way, in-out barrier. It absorbs nutrients,
oxygen, and other useful substances as part of nutrition and
respiration, and removes wastes and unwanted substances as
part of excretion. These dual functions are especially important in
various types of worms, such as segmented worms,
atworms, ukes, and tapeworms, because
many of them lack a well-developed
respiratory or circulatory system. As a
result, they rely on their selectively
permeable skin to absorb oxygen and
remove carbon dioxide and wastes in direct
Parasitic tapeworms have no mouth
exchange with the environment. In the sea, arrow
or digestive system. Instead, their very
worms have no excretory system and expel their
thin skin absorbs nutrients from the
body wastes through the skin.
semidigested gut contents of their hosts.

Slippery skin
In aquatic animals a specialized function of skinin addition to its
many standard functionsis to minimize drag, turbulence, and
eddying for fast, energy-efcient movement through the water. For
example, dolphin skin sheds tiny particles of skin into the water
owing past, which greatly reduces drag. The sloughing skin layer
replaces itself every two to four hours. In addition, the skin has
microscopic ridges that hold a thin layer of water between them.
So, in effect, the dolphins outer skin layer consists partly of water
as it swims, thereby reducing drag even more, as water slips past
water. A third adaptation is the dolphins very exible skin. As the
dolphin swims, the skin distorts, bending and rippling
into contours that yield least resistance to the ow of
water, which again minimizes drag.

The skin of the African
elephant is up to 2 in
(5 cm) thick in places
and is very tough. In
some areas, such as
the tip of the trunk, the
skin is much thinner
and is highly sensitive.

A dolphins skin
adapations mean
that drag is reduced
more than 100-fold
compared to standard
mammalian skin, which
enables bottlenose
dolphins to swim faster
than 19 mph (30 kph).

Sea squirts have a slippery,
leathery skin that forms a
tough, overall covering.
Unlike most animals, the
skin is composed of a type
of carbohydrate (known as
tunicin) that is simlar to the
cellulose found in plant cells.

103 SKIN



Most scales are small, platelike outgrowths from an animals skin or body
covering. They provide protection while allowing exibility, and partly determine
visual appearance. Various behaviors, including parasite removal, self defense,
camouage, and courtship are linked to the structure and appearance of scales.

Origins of scales

Vertebrate scales

Several groups of animals have scales or scalelike

coverings, from worms and certain insects to shes,
reptiles, and birds. There are also scaly-looking
mammals such as armadillos and pangolins. In
most of these animal groups, scales have evolved
independently. Others show an evolutionary
progression; for example, the scaly legs and feet
of birds may be derived from their reptile ancestors,
specically small, meat-eating dinosaurs.

Although some birds have scaled legs and feet,

and mammals are represented by the unrelated
armadillos and pangolins, the vast majority of
scaled vertebrates are shes and reptiles.
There are several types of sh scale. Cartilaginous
shes, such as sharks and rays, have teethlike
placoid scales. Each consists of a plate of enamel
embedded in the skin, a main body of dentine,
and a pointed cap or spine of enamel. Ganoid
scales are found in primitive ray-nned shes such
as sturgeons. These also have dentine, as well as
enamel-like ganoine surface layers. In the gar,
diamond-shaped ganoid scales t closely to form a
complete covering. In sturgeons the scales are large,
platelike, and thickened with bone, forming scutes.
Fishes that are more advanced in evolutionary terms

Invertebrate scales
Among invertebrates, scales are found in a group of
segmented worms called scaleworms. They have 12 or
more pairs of leathery scales or elytra that overlap like
roof tiles. These form a protective outer covering for
these stout, free-swimming, predatory worms, but also
form channels for water currents that help them breathe.
Butteries and moths form the major insect group
Lepidoptera (the name means scaly winged).
Their wing scales are attened versions of
Flaplike scales
less than
0.1 mm long
cover the wings
of butterflies
and moths.


insect hairs called setae. Numbering thousands per

wing, these tiny scales are arranged in rows, and
their layout and colors produce the wings overall
appearance. They also aid air ow, which improves
aerodynamics and helps maintain body temperature.
The males of some species have scent scales on the
forewings that release pheromones to attract females.




From microscopic flakes to cumbersome
plates of armor, the variety of scale types in
vertebrates reflects the diversity in form and
behavior of the animals that bear them.
A major advantage of a scaly
covering is that it combines
protection with flexibility for
movement. The junctions between
the scales are made of proteins
such as collagen or keratin and
are tough yet bendable.

have thin, smooth, overlapping cycloid scales, as in

cod, and rough-surfaced ctenoid scales, as in perches.
The scales consist of a calcium-rich base with collagen
bres, similar to bone, covered by a thin outer layer.
Among reptiles, lizards and snakes have scales
made of horny, aplike, overlapping extensions of
the outer skin, made chiey of keratin (the tough
protein that forms bird scales and feathers, and
mammalian skin, horns, hoofs, claws, and nails). These
contrast with the solid, fused, bone-reinforced scutes
embedded in the outer skin, as seen in crocodiles and
alligators, and especially in turtles and tortoises.

old skin and


new scales
revealed beneath

Most lizards and snakes shed their skin and scales
several times yearly. The epidermis and scales
come away to reveal newer, larger scales beneath.

HOW FISH SCALES GROW Most sh scales originate in the
dermis, the inner of the skins two layers. Although the materials
they contain differ, most scales develop in a similar way. The
placoid scales of sharks (shown here) begin as small clumps of
dermal cells, which form tiny mounds. As these mounds grow



mound has


taller, they tilt backward and the cells on the upper surface secrete
a hard dentine layer with a cap of even harder enamel above it.
This produces a tough and abrasive skin surface. These scales
do not enlarge as the shark growsit will grow more scales to
cover the expanding body surface.

row of enamelsecreting cells


HOW REPTILE SCALES GROW In lizards and snakes,

scales begin as pointed mounds in both the dermis
(inner) and epidermis (outer) skin layers. As the dermis
ages, it leaves the epidermal scale, which is hardened
with a protein called keratin.

hardened scale









Scale variety
In more primitive fishes, scales
develop as thickened areas of
skin that are arranged side
by side, limiting flexibility.
Advanced fish have much
thinner, lighter scales growing
from pockets of skin. These
have free edges that can be
lifted or tilted, enabling
flexibility and free movement.

Most lizards and snakes have
overlapping scales arranged in
diagonal rows. There are also
various modified scales forming
flaps, spines, frills, and spikes. In
addition to providing protection,
scales help retain moisture. By
emphasizing skin color, they play
a role in defense, courtship, and
territorial displays.
The bearded dragon has rows of spiked scales along the sides of its
body, which it uses for defensive displays.

Coelacanths have modied overlapping cosmoid scales, with tiny

toothlike spines, or denticles, on each exposed portion.

The gars thick scales have a layer

of very hard ganoine.

The marine iguana has differently shaped scales forming knobs,

spikes, and cones, used especially in visual displays.

In many sharks, pointed placoid scales produce a rough, abrasive

surface. A sharks teeth are actually larger versions of its placoid scales.
A rattlesnakes rattle is
made of unshed scales.

Ctenoid scales, like those of the garibaldi, have a surface with

minute ridges or toothlike projections.

The Cape legless skink burrows Reptile scales are mostly transparent,
allowing the skin pigment to be displayed.
with a strong snout scale.

Cycloid scales have smooth surfaces

and edges, as in the trout.

Wide scales on a snakes underside

aid in locomotion.

A snakes eye is covered by a single clear scale,

the spectacle or brille.

Most sh scales are transparent, allowing skin

color to show through.

Modied scales form many kinds of accessory skin

structures, such as the porcupine shs spines.

Scales thickened with bone, forming scutes, offer

extra protection to the pineapple sh.

The horned vipers horns are scales that may work as

camouage by breaking up the outline of its head.

The skin of crocodiles and alligators is covered with

nonoverlapping scales embedded with bony scutes.

bOdy cOverIngs 106


Protection and conditioning

Of all living animals, only birds have feathers, which are

made from the tough, fibrous protein keratin. In addition
to being used for flying or swimming, feathers have many
other functions, including protection, insulation,
waterproofing, camouflage, and display.

Feather types
Most birds have different types of feathers on different parts of the body. The
main flight feathers (sometimes called remiges) are the primary feathers; they
are located toward the wing tips and can spread out like vanes of a fan. On
the inner wing are the secondary flight feathers. Tail feathers, like the
primaries, can also move and spread. Contour feathers form
a streamlined surface over the body and
where the wings join the chest. Beneath the
contour feathers are fluffy down feathers,
which trap air to provide insulation.
fluffy to smooth
The contour feathers
form a smooth surface
over the body but may
have downy, flexible
plumes at their bases.
These plumes do not
interlock and provide
insulation and
cushioning. The flight
feathers have little or
no downy base.

secondary flight


primary flight

feather structure
Feathers grow from the epidermis of the
skin and typically have a long, hollow shaft
(the quill, or rachis), supporting a large flat
surface, the vane. The vane is made up of
small parallel strips called barbs, which
are like the teeth of a comb. The barbs
bear even smaller branches, barbules,
some of which have interlocking hooks.
As a bird preens, it arranges the barbs
into neat rows to form an air-proof surface,
which is essential for efficient flight and
also provides insulation.

barbule with
no hooks

Flight control
The primary feathers at the wing tip can be fanned out and twisted to
adjust their resistance to airflow. This action provides the birds main form
of aerial control, allowing it to slow down, rise, descend, and bank. At
slower speeds the tail is also important, used as a rudder for maneuvering
and as a fanned-out air brake for landing. The secondary feathers are less
adjustable, but form an arched, airfoil surface that
generates lift as it moves through the air. A small tuft
of feathers at the leading edge, the alula or bastard
wing, can be used for low-speed maneuvering.

primary flight
secondary flight

inner wing

wing parts
The flight feathers are long and strong. They
are blended into the wing structure and body
by smaller covert feathers on the inner wing,
as shown on the wing of this shoveler.

The outer feathers give physical protection, while the down feathers
beneath provide cushioning. Feathers also provide waterproofing,
which is particularly important for birds that spend time in water, such
as seabirds, wildfowl, and waders. As a bird preens, it cleans and tidies
the feathers, removes dirt and parasites, and arranges the barbs into neat
rows. The bird also spreads oils from its skin oil glands over the feathers,
so that they resist water absorption and shed moisture
easily. Many birds molt twice yearly and replace
the shed feathers with new plumage.
This gets rid of damaged feathers and
also alters the birds appearance, for
example, for a spring breeding display
or fall camouflage.
In addition to feeding, nest making, and defense, a
birds bill is used for cleaning and combing through
the feathers, arranging them correctly, and
waterproofing them, as this roseate spoonbill is doing.
slow flier
This waxwing demonstrates control
at slow speed as its primary wing
feathers and tail feathers fan
out to increase air resistance
and the tuftlike alula
feathers are raised to
give better lift and
prevent stalling.

Fur, hair, and bristles

True fur or hair is found only in mammals. It is composed
of strands of the brous protein alpha-keratin, which is
made by specialized structures in the skins epidermis.
Fur, hair, and bristles also occur in other animals but
they are made of different materials.

long, coarse
hairs of outer
guard coat

short, dense
hairs of inner

Seal fur has the two layers typical of
mammalian fur. Long, coarse hairs
form the protective outer guard coat
and shorter, denser hairs constitute
the insulating undercoat.

Mammalian fur has a wide

range of different functions:
physical protection and defense;
insulation of the warm-blooded
body; camouage or brightcolored display in some species
or at certain times of the year;
waterproong in semiaquatic mammals;
and various sensory functions. Each hair
grows from a hair follicle, a small pocketlike pit
in the dermis. Insulation is improved by ufng up
the hairs using the tiny arrector pili muscle attached to the
base of each hair follicle. This same action is used in aggressive
or defensive behavior to make the mammal look larger, as when a
dog or wolf raises its hackles.



Many animals with attractively colored or
patterned fur coats have become endangered
from being hunted for their pelts. Species
include big and medium cats, for example,
tigers, ocelots, and jungle cats, as well as
foxes, minks, chinchillas, coypu (nutria),
beavers, and fur seals. Captive breeding
on fur farms and restriction of trade
in furs and skins by the CITES
agreement have reduced the
need for wild kills.

The bristles of woolly caterpillars like this sycamore
moth larva are made of chitin rather than keratin. In
many species, the bristles break easily when touched
to release noxious chemicals as a defense mechanism.


Fur insulates against cold, as in seal
pups (top), and against heat, as in
bactrian camels that live in the hot
desert of Central Asia (above).

Touch and sensation


Hair or fur shafts are almost entirely dead, except at the root in the base
of the follicle, where cells accumulate to increase length. The cells quickly
ll with keratin, cement together, and die to form a rod or tube shape as
they move up the follicle with the growing shaft. The nerve endings wrapped
around the follicle are sensitive to a hairs movement when it is tilted or bent.
Mammals use this sense to detect direct physical contact, and they also
use the hair movements to gauge wind or water currents. Some
animals have whiskers, extra-large hairs with follicles that are
specialized for
touch. Whiskers
are especially
important in
nocturnal animals
such as cats and
rats, where they
extend the width
of the head, thereby
enabling the animal
to feel its way and
assess if gaps are
Aquatic mammals, such as this beaver, and also otters, seals,
big enough to pass
sea lions, and walruses, tend to have many whiskers. They
through, even
enable the animal to feel its way in murky water, especially
at night, and to locate food by touch.
in darkness.

A mammal relies on its fur for survival, so it must be kept

in good condition. Grooming, carried out with the teeth,
claws, or nails, gets rid of dirt, mud, pests, such as lice
and eas, and tangles in the fur. It also spreads natural
skin oils (sebum) from the sebaceous glands of the hair
follicles, to keep the hairs smooth, pliable, and waterresistant. In some groups of mammals, mutual
grooming occurs not only for hygiene but also for
social reasons. It is part of the parental care of offspring
and also a way of establishing a
close bond between a breeding
pair or rank in a hierarchy.
Usually the animal groomed
the most has the higher
rank or is dominant and
the lower-ranking or
submissive animal does
most of the grooming.

Some primates, such as
this diademed sifaka,
have a row of forwardpointing teeth in the
lower jaw that are used
as a tooth comb for

Chimpanzees show dominance and submissiveness
by the relative amounts of time spent grooming
each other. Grooming also strengthens alliances
between individuals who regularly forage together.

Jellyfish like the sea nettle seem delicate and
relatively simple, with only a few organs. But, like
all animals, they need to maintain their internal
conditions of dissolved nutrients, salts, minerals,
and other substances; otherwise, their
physiological processes would grind to a halt.


All animal bodies carry out similar inner processesbreaking down food for energy
and nutrients, obtaining oxygen for energy release, getting rid of waste, and
coordinating internal parts so they work together properly. In some animals, these
processes occur in all tissues; others have complex body systems for each.

In, between, and out

The ins of living involve taking in essentials such as nutrientswhich
provide energy and raw materials for growth, maintenance, and repair
and oxygen, which is required chiey to release energy for useful work.
The outs involve removing waste, leftovers, and potential toxins from
the body. Between the ins and outs, it is vital to maintain a suitable
environment within the body, in terms of the amount and concentration
of all kinds of substances, from water to complex organic chemicals. This
is the concept of homeostasisthe constancy of the internal environment.
It involves physiologythe functional, or biochemical, side of how living
things work, usually at molecular level. Physiology complements anatomy,
which is the structure of the body and what it is made of. The collective
name for all the thousands
of biochemical processes
in the body is metabolism.
All of these processes are
under the control of chemicals
known as enzymes, each
of which regulates a
particular reaction.

A flatworm has a nervous system for
coordination, and a digestive system.
But it has no proper circulation with a
heart and flowing fluid, and no
specialized respiratory system.
Crayfish, like other arthropods,
possess all the major body systems
of vertebrates such as mammals, but
generally in a more simplified form.

In mammals such as the wolf, each
system is composed of several main
parts, called organs, which may be
close together or widely separated
around the body.


Life chemicals and processes

In the scorching desert, air temperatures may
rise above 120 F (50 C). The ostrichs body
is fine-tuned to stay at about 18 F (10 C)
lessany higher disrupts its internal
chemistry. So the ostrich uses anatomy,
physiology, and behavior to stay cool.
As temperatures fall to -22 F (-30 C) or less,
the Arctic hare would be in danger of freezing
solid. It fluffs its fur to trap insulating air, and
shelters from the biting wind, so that its body
can remain at a constant warm temperature.

Key to major systems

Food is vital for all animals. This is dealt with by the digestive system,
which breaks up food with enzymes. Eventually, the pieces of food are
small enough for the bodys tissues to absorb. In simple animals, the
nutrient molecules may just drift through the cells and tissues. In more
complex animals, there is a circulatory system that propels a uid, such
as blood, around all body parts to deliver the nutrients. Likewise, oxygen
may simply be absorbed at the body surface and pass into the tissues,
or it may be taken in through specialized parts, such as the lungs or gills
of the respiratory system, and then passed to the circulation for bodywide
delivery. In a similar way, waste may diffuse outward to the body surface,
or it may be collected and disposed of by an excretory or urinary system.
The immune system protects an animal from germs and disease.
Coordinating all these systems are the nervous and hormonal systems.

The energy sources that power most life processes are sugars, especially
glucose obtained by digestion. In cellular respiration, glucose is broken
apart to release its chemical energy. This is then transferred to energy
carrier molecules known as adenosine triphosphates (ATPs). In complex
animals, oxygen is brought into the body by the respiratory system
and then distributed by the blood. Each overall
reaction begins with one molecule of glucose and
six of oxygen, and yields six molecules each of the
waste product carbon dioxide and water, plus
released energy.
glucose molecule

oxygen combines
with glucose

oxygen diffuses
out of blood

six water

Animals that hold their
breath for a time, like sea
snakes, can alter their
cellular respiration so it is
anaerobicit does not need
oxygenbut only for a while.

six carbon
tissue cell

into blood


six oxygen

Red blood cells bring continual
supplies of fresh oxygen and
glucose. These pass through
the thin walls of the smallest
blood vessels, the capillaries,
into cells. The carbon dioxide
produced by cellular respiration
diffuses into the blood and is
removed by bodily respiration.

body sysTems 110

The term breathing is usually applied to the physical
movements of inhaling and exhaling. It partly overlaps
with the broader term respiration, which can refer
to the overall process of taking in oxygen or using it
to release the energy from glucose and similar
nutrients inside cells.

Breathable skin
Skin can be breathable in the sense that it is a gas-exchange surface
through which oxygen is absorbed into the body from the environment,
as carbon dioxide passes the other way. This is known as cutaneous
respiration. The skin must be thin to present a minimal barrier to the
diffusion of gas. This type of breathing occurs in aquatic animals, where
oxygen is dissolved in the water. The amount of dissolved oxygen rises as
the water circulates or becomes colder. So cool, fast-flowing streams
have abundant dissolved oxygen, while tropical swamps have much less.
Swamp-dwelling fish can absorb oxygen through their skin as well as their
gills, and some, like lungfish, gulp oxygen into their lungs.
cASe STudy

lungless frog
Most frogs respire through their skin and
also have lungs to breathe air. The recently
discovered Bornean lungless frog, the firstknown frog species without lungs, shows
that under suitable conditions, enough oxygen
can be absorbed through the skin alone
in this case, in a habitat of cool streams.
Amphibians evolved true lungs more
than 300 million years ago. The lungless
frog and several species of lungless
salamanders have reversed this trend.

Invertebrate breathing systems

Some terrestrial invertebrates, especially insects, have a respiratory network
of air tubes (trachea) branching throughout the body. The tubes open at
holes in the body covering called spiracles. Air movement through the
spiracles, into and out of the trachea, is much more limited than the forced
airflow in true lungs. It occurs mainly when the insect moves, making the
trachea compress and stretch, or from air currents. However, even in still
air, oxygen can diffuse to areas inside the trachea where there is less oxygen,
while carbon dioxide does the reverse and is removed. Most spiders have
book lungs in a chamber in the base of the abdomen. Book lungs consist
of many thin, leaflike structures into which oxygen can easily diffuse.

insect spiracles
Spiracles are seen clearly along the sides of
insect larvae such as this sphinx moth
caterpillar. They may be incorporated into
coloration, perhaps as a form of disruptive
camouflage that breaks up the body outline.
hole in the wall
Each spiracle can become smaller by contraction of
the closure muscle around its opening. This reduces
loss of moisture from inside the body as vapor in dry
conditions. Opening up increases the flow of oxygen
to the muscles, so the insect can move more.

plenty of surface
Flatworms, like this
marine turbellarian, lack
a circulatory system to
distribute oxygen. They
also have no specialized
respiratory parts such as
gills. The leaflike body
shape allows tissues to
be a minimal distance
from the skin surface, to
receive dissolved oxygen
as easily as possible.

external gills
Amphibian larvae, like
this newt, have external
gills on the sides of the
head. These are delicate
and easily damaged
but regrow well.

111 breathing

Breathing with gills

feathery gills

Gills are body parts specialized for

gas exchange in water. Their structure
consists of many branching surfaces
with a plentiful blood supply, to present
the greatest possible area for absorbing
oxygen and getting rid of carbon dioxide.
Gills of various kinds are found in a wide range
of aquatic animals. They form frilly tufts on the
backs of sea slugs, and fin- or flap-like tail
appendages on aquatic insect larvae such as
damselfly and mayfly nymphs. Fish gills are on several bony or
cartilaginous arches on the sides of the head. In all cases, the
gills must be exposed to flowing water, to bring continuing
supplies of dissolved oxygen. The flow also takes away carbon
dioxide and other unwanted substances, such as salt and
ammonia. This occurs in fishes and some amphibians, where
gills have become organs of excretion as well as respiration.

oral valve

direction of water
gill arch, attachment
point for filaments


internal gills
Fish gills are composed of hairlike or
feathery filaments protected within gill
chambers. In most fishes, water flows in
through the mouth, over the filaments
where oxygen is taken into the blood
inside themand out through the gill
slits (left). The oxygenating blood gives
the gills a strong red color, as in the map
pufferfish (above).

Breathing with lungs

Lungs are organs specialized to absorb oxygen from air and
remove carbon dioxide into air. Most vertebrates except
most fishes have lungs (some fishes, such as lung fishes,
take in oxygen from swallowed air). Typical vertebrate
lungs are paired in the chest, on either side of the heart.
Their branching airways connect to the trachea (windpipe),
pharynx (throat), and mouth, forming a passage along
which air moves. The flow is caused by respiratory
muscles in the chest and the sheetlike diaphragm between
the chest and abdomen. These muscles contract to expand
the lungs and suck in fresh air
one-way lungs
during inhalation, then relax
Mammal lungs are dead ends in
so the stale air pushes out.
that air flows in, then back out. Birds
Lungs contain millions of tiny
have expandable air sacs that draw
bubblelike alveoli surrounded air down the windpipe and right
through the lungs. This allows more
by blood capillaries, which
oxygen to be absorbed, for the birds
energy-hungry flight muscles.
absorb oxygen from the air.

cervical air

air sac


air sac

abdominal air

posterior thoracic
air sacs

land crab
The coconut or robber
crab has a combined
gill-lung known as the
branchiostegal organ in
its rear abdomen. If
moistened with sea
water, this can take in
oxygen from air for long
periods, allowing the crab
to move considerable
distances on land.

case sTudy

record dive





Depth (feet)

Depth (meters)

holding breath
Most seals, such as this
Antarctic fur seal, for example,
and sea lions can hold their
breath for at least several
minutes, and some for over
one hour. The body uses
oxygen stored not only in the
lungs, but in the blood and
muscles, by the pigments
hemoglobin and myoglobin.

Cuviers beaked whale has been

tracked to almost 6,200 ft (1,900 m)
below the surface on a single dive
of 85 minutes. Compared to land
mammals, whale blood contains
very high amounts of the oxygenloving pigment hemoglobin, but
large quantities of a similar
pigment, myoglobin, are found in
the muscles. These store plenty
of oxygen for diving. Blood vessels
to less important body parts like
the intestines constrict on the dive,
saving oxygen usage, while the
vessels to the muscles, heart, and
brain stay open for a plentiful flow.






time (minutes)



assisted breathing
As large animals such as the pronghorn run, the
continual acceleration-deceleration of each stride
makes the abdominal contents move to and fro
within the body. This aids normal breathing by
alternately compressing and stretching the chest.


In complex animals, the circulatory system sends blood around the
body, through a network of tubes or vessels, pumped by the heart. The
circulatory systems of other animals use a different uid, or have few
vessels, or no heartand sometimes they have all these variations.

Types of circulation

The throbbing
heart of a waterflea (a small
is visible through
its body wall.

A dedicated circulation system is much more efcient than

simple diffusion, where substances ow at random through
tissues and cells. The two main types of circulation are open
and closed. In the former, found in invertebrates such as
insects, the circulating uid is usually called hemolymph. For
part of its journey, it permeates and oozes through the general
body cavity, the hemocoel or celom, unconstrained by
vessels, before returning to the heart(s). In the closed system,
as seen in most vertebrates, the circulating mediumblood
is within vessels for all of its journey. It exchanges nutrients,
oxygen, waste, and other substances through their walls.

Red blood has the iron-based
pigment hemoglobin carried
in red cells (above). Some
crustaceans and mollusks have
copper-containing pigments,
like hemocyanin, so the
hemolymph is green or blue,
as in a whelk (left).

Blood, lymph, and hemolymph


The blood of vertebrates, and the hemolymph of

invertebrates, carries out dozens of functions. It
transports nutrients and oxygen to the cells, and
gathers unwanted by-products for excretion. It carries
hormones to coordinate inner processes. It gets sticky or
clots to seal wounds and leaks. In warm-blooded
animalsmainly mammals and birdsit distributes
heat around the body. In a typical vertebrate, about half
the blood is a pale uid (plasma) containing hundreds
of dissolved substances. Most of the rest is red blood
cells (erythrocytes), which hold onto oxygen or carbon
dioxide. Lymph is a circulating uid that has important
roles in the immune system. It is carried inside
vessels called lymphatics. It has no pump but
oozes slowly, massaged by body movements.


Immune system
The immune system has major roles in resisting
disease and ghting illness. It involves various
categories of white cells. Those called macrophages
hunt down, engulf, and eat microbes such as
bacteria. There are also various kinds of lymphocytes.
Some are able to recognize microbes and other
alien items. They instruct other types of
lymphocytes to produce substances known as
antibodies, which stick onto the microbes and
disable or kill them. In vertebrates, all of these
cells travel in the blood and the lymph.

abdominal vein

A glass frog reveals its heart at the front of the
chest cavity. The thick muscular walls obscure
the red blood inside, but blood can be seen in the
main vessels leading away.

Animals sometimes show signs of
infection, as in the case of this elephant
seal. Mucus is produced by the nasal lining
as the white cells of the immune system
attack invading germs there. Some seals
suffer from phocine distemper, related to
canine distemper, which affects dogs.

Consuming food for internal digestion and absorption distinguishes
most members of the animal kingdom from those in other
major groups of organisms, such as plants and fungi. Food
provides raw materials for growth and repair, healthgiving substances like vitamins, and the energy for life.

Digestive systems
After an animal captures food it is taken in through the oral opening, or
mouth, into the digestive tract. In simple animals, this is a hollow chamber
or branching system with just one opening, so the undigested waste
comes out the same way and the mouth functions as the anus. In complex
animals, the digestive tract is a long tube or convoluted passageway with
the anus or cloaca at the other end. General names are given to sections
of the tract in various animals. After the mouth is the gullet or esophagus,
perhaps leading to a crop which is specialized for storage. The stomach
is the main digestion site, the intestine the chief area for absorbing
nutrients, and the rectum, or large bowel, stores waste until it is expelled.


mouth or
large intestine



small intestine

Cnidarians such as coral polyps (above), jellyfish,
and anemones have a single digestive opening
that is the mouth when eating and the anus when
expelling leftovers. Nutrients diffuse through the
gastrodermis (gut lining) into the tissues.


In some invertebrates and all vertebrates, the gut
is a through-tube. Birds, some fishes, and some
invertebrates have a gizzarda muscular grinding
chamber following the proventriculus (the stomach
area) that secretes digestive enzymes.

In animals such as flies, spiders, and starfish,
early digestion is outside the body. The mouth
pours digestive juices onto the food, which
turns it into a soup that is sucked up.

Specialist diets
Some animals are omnivorous, with a digestive system that can deal with
many kinds of foods. Others are adapted to survive on a narrow range of
foods, especially those that are very low-nutrient, or distasteful, or contain
chemicals that are toxic to other animals. This means the specialist feeder
faces little competition for its meals, but also
limits its range to the geographical distribution
of its foodstuffs, and its survival to that of the
food. Some specializations are physical, for
example, the manipulation of bamboo shoots
by the giant panda. Others are biochemical,
where a unique enzyme allows an animal to
deal with a particular food part or a toxin.

Food breakdown and disposal

Digestion involves breaking food into tinier pieces, until they are small
enough for absorption. Physical digestion includes crushing in the mouth
and mashing in the stomach or gizzard. In chemical digestion, juices
containing enzymes are secreted onto the food by the gut lining. Different
enzymes attack different dietary constituentsproteases split apart proteins
in meaty food, while lipases break down oils and fats in fatty food. This
process may take months, as when a python digests a wild pig. In most
cases, undigested material passes out through the anus as droppings or
feces, but some animals bring up or regurgitate leftovers via the mouth.
hair and

The golden bamboo lemurs
staple diet of giant bamboo
(left) contains levels of cyanide
that would kill most other
mammals, while the creosote
bush katydid (above) is not put
off by the unpalatable taste of
an acid that is found in
creosote bush leaves.

vole leg

Worm casts, like that of the lugworm, consist of sand

and mud particles that have passed through the gut.

Owl pellets are regurgitated and contain undigestible

bits of prey such as bones, teeth, fur, and beaks.

body systems 114

Fluids and temperature control

the control of an animals bodily internal environment includes regulating the concentrations
of hundreds of salts, minerals, and other substances. Called osmoregulation, this involves
the delicate processes of water balance. the body temperature must also be maintained
within suitable limits so that biochemical reactions can take place efficiently.

Water and salt balance

Living on land, in freshwater, and in seawater pose different problems.
Land animals tend to lose water as vapor from permeable body coverings,
from moist respiratory surfaces, and in excreted urine and droppings. This
must be replaced by drinking, by water contained in foods, and by water
made in the body by metabolic processes. In freshwater, an animals
internal environment has a relatively high concentration of salts compared
to its surroundings. Water tends to diffuse into the body and so must be
removedin urine and by being actively pumped out.
In saltwater, the opposite may occur, so water
must be prevented from leaving the body.

sleeping bag
The desert-dwelling
water-holding frog buries
itself in moist soil during
drought. It forms a
watertight cocoon from
shed layers of skin,
storing water under its
skin and as dilute urine in
its large bladder. The frog
remains inside its fluid
sleeping bag until wet
weather returns.

metabolic water released by

digesting food

moisture in food


water in

moisture in droppings


moisture lost from skin

and in breath


water out

desert dweller
Desert animals such as the banner-tailed
kangaroo rat have adapted to produce very
concentrated urine, which reduces water loss.
This kangaroo rat is nocturnal, and it seals its
burrow by day, which traps moisture in its breath.

water balance in a kangaroo rat

There is little moisture in the kangaroo rats main
diet of seeds. But water is actually made in the
body by metabolic processes such as cellular
respiration. As well as concentrated urine, the
droppings are also dry, to conserve moisture.

Waste disposal

useful droppings
A birds urine, produced
by its kidneys, is combined
with digestive wastes
from its intestine, and
leaves the cloaca as
semiliquid droppings.
Thick accumulations
known as guano from
seabirds such as the bluefooted booby are collected
for fertilizers and mineral

Filtering blood, hemolymph, or other body fluids is a

common method of regulating water, salts, and toxins. In
vertebrates, the main organs involved are the two kidneys.
They remove waste products dissolved in water through
microscopic filters known as nephrons. Then the required
amount of water is reabsorbed into the bloodstream to
maintain water balance. In mammals, the resulting fluid,
urine, is stored in the bladder until it can be expelled. Urine
contains various hormones and similar substances, which
often have distinctive scents, so it has a secondary role as
a means of communication, for example, as a sign of
readiness to breed. Invertebrates have similar methods
of excretion, involving filtering body fluids such as
hemolymph, but the main
organs have different
structures. Worms or
flatworms possess
nephridia, while the
excretory system of an
insect is based on
malpighian tubules.
useful urine
Behavioral aspects of waste disposal
include the use of urine to scentmark territories and warn away
intruders of the same species, as
seen in lions, rhinoceroses, and
many other mammals.

cool ears
Elephants have large ears, not
only for acute hearing, but to help
control their body temperature.
The ears have a plentiful blood
supply, and when the body
temperature rises, the ears flap
back and forth to work like
radiators and lose heat to the
surrounding air. Animals with
similar adaptations for hot
climates include jackrabbits and
the fennec fox.

Mammals and birds maintain a constant high body temperature by burning

energy-containing nutrientsto release heat. Keeping body temperature
constant is termed homeothermy, popularly known as being warmblooded, and generating warmth in this way is termed endothermy. Such
animals can stay active even in cold conditions, but the process requires
energy, which must come from increased food consumption. Most other
animals are ectothermic, meaning that warmth for their bodies comes from
outside. A popular term for this is cold-blooded. However, a reptile in a
scorching desert may have a warmer body than a mammal next to it. Also,
ectothermy does not mean having no
controlby using behavior
such as resting in the shade
to cool down, an ectotherm
can alter its temperature.

conserving warmth
Mammals and birds in cold places,
such as penguins in the Antarctic,
have a specialized blood flow called
the countercurrent mechanism to
conserve body heat. Extremities
such as feet and flippers tend to
cool fastest. Warm blood flowing
from the body core to the extremity
passes close to cool blood returning
from it, and transfers some heat to
warm this blood, while itself cooling.
The extremity is kept colder and so
loses heat at a slower rate than if
warm blood circulated through it.
warming up
In the early morning, a southern
rock agama seeks out dark rocks
to warm itself. These rocks have
held the suns heat from the
previous day and are also
absorbing more solar heat in
the new dawn.

direction of
blood flow
warm blood
returns to body
warm blood
from body
transfers heat
to cold blood
cold blood
circulates in
penguins foot

Hibernation and torpor

True hibernation is limited to certain mammals such as bats,
dormice and other rodents, insectivores including hedgehogs,
and some lemurs. It is a strategy to survive adverse conditions,
usually winter, by shutting down the bodys activity and
metabolic processes to save energy. Heart and breathing rates
fall to a fraction of their usual levels, and body temperature
drops to a few degrees above freezing. To prepare, the animal
feeds well to lay down reserves of food as body fat, then finds
a sheltered, safe place. Once in hibernation, it is unable to
rouse quickly. Torpor is a less extreme, short-term
slowing of body processes, usually just for a few
hours. Some small bats and hummingbirds
enter torpor overnight to survive the cold.

fat reserves
The fat-tailed lemur stores
food as fat in its tail to survive
Madagascars dry season,
which it spends in a state of
torpor in a hollow in a tree.

cASe STudy

lizard activity patterns











time of day

Air temperature

Sheltering to avoid cold

Normal activity

Lizards body temperature


Sheltering to avoid heat

temperature f

temperature C

Terrestrial ectotherms, such as

lizards, have an array of behaviors to
help them warm up by day, and then
keep their body temperature
relatively constant so that they can
remain active. They move from
shelter to sunshine and bask on dark
rocks, which soak up the suns heat
better than light-colored ones. To
cool down, they seek out shade or a
breeze, gape the mouth to breathe
out warm air, or enter a burrow.

safe cave
Hibernating bats, such as
the whiskered bat, choose
sites that are safe from
predators, and which also
have constant conditions
and do not freeze, for
example, deep inside
a cave.

115 fluids and temperature control

Behavior and temperature control


Brains, nerves, and hormones

The nervous and hormonal systems of animals are vital for the control and
coordination of internal body parts, ensuring they work together effectively. These
systems also control the whole animal as it sees, hears, and otherwise senses its
surroundings, moves around, selects a range of behaviors according
to circumstances, and prepares to molt or breed.

Most animals possess a system of nerves that branch into all
body parts and come together at one site, known as the brain,
or at several locations, where they form ganglia. The nervous
system uses tiny pulses of electricity, or nerve signals, and is
concerned with the whole animal sensing and reacting to the
environment, instincts, memory, and learning. Its basic
components are some of the most specialized of all cells
neurons, or nerve cells. They have thin
branches that carry nerve signals
at speeds in excess of 330 ft/s
(100 m/s) in some species. The
branches almost touch those of other
neurons, but are separated by tiny gaps
known as synapses, which the nerve
signals cross in the form of chemicals
called neurotransmitters released by the
neurons. Not all animals have a nervous
system and brain. Sponges lack any
nerves and jellysh are brainless, with
only a simple nerve net.

As part of the nervous system, millions of cells
give physical and nutritional support to neurons,
without carrying any nerve signals themselves.
Astrocytes (left) are named after their starlike
shape, and pass nutrients to neurons.
Oligodendrocytes (above) form a type of
living scaffolding to hold the neurons firmly.


cell body

A typical neuron has a rounded cell body
with short, thin branches called dendrites.
These gather signals from other neurons
and process themthe resulting signals
travel along a thicker, longer projection,
the axon (nerve fiber), to the terminal,
which links to other neurons. Some
axons have wrappings of fatty insulation
(myelin) made by Schwann cells.

Schwann cell

Many short, thick nerves
run from the sense organs
of smell, vision, taste,
and touch to a sharks brain
in its skull. In sharks and
rays, thin nerves also run
from the tiny electrosensing
pits, ampullae of Lorenzini,
scattered over the snout.

node of Ranvier

axon terminal

Invertebrate nervous systems

Invertebrates show a range of nervous system designs,
from simple nerve nets to centralized networks, which
have lumps of neurons called ganglia or a single
brain. These structures contain concentrations of neuron
cell bodies with many short, interconnected dendrites
and axons. This makes the exchange and processing
of information more efcient than in a diffuse network.


Sensory nerves bring incoming

signals from the sense organs.
This information is analyzed in the
brain or ganglia and appropriate signals
are then sent out along motor nerves to
muscles, which effect movements and
behavior, and to body parts such as glands,
telling them to release their chemical secretions.


buccal mass



ventral nerve cord



nerve net





The nerve net of cnidarians such as the
hydra, a tiny pond animal, consists of
nerve fibers connected in a simple netlike
fashion. Arthropods such as flies and
other insects have a frontal brain, ganglia
at various sites, and a ventral nerve cord
along the base of the body. Mollusk
nervous systems, shown here in the
snail, have several ganglia linked by
thick tracts of nerve fibers that carry
signals at very fast speeds.

The vertebrate brain, colored green in this
image, is well protected inside the cranium,
a bony chamber at the rear of the skull. Its
wrinkled surface provides a large area for
billions of neurons.

The typical layout of a vertebrates central nervous system consists of a brain

and spinal cord, and branches from these to all body parts, forming the
peripheral nervous system. The presence of a spinal column made of vertebra
(backbones), which support and protect the spinal cord, is a characteristic of
vertebrates. Different lobes, centers, and other parts of the brain deal with
specic functions. For example, the optic lobes receive nerve signals from the
eyes, while the olfactory lobes process information about smell from the nose,
and the motor centers organize nerve signals going out to the muscles for
movement. There is also an autonomic nervous system, partly with its own
nerves such as sympathetic ganglia chains, and partly using nerve bers from
the other systems. This system is concerned with the automatic running of
essential actions inside the body, such as breathing and the passage of food
through the gut. The sympathetic part of the autonomic system makes body
parts more active and ready to cope with stress, while the parasympathetic
part restores calm and normal working.

Many types of sensory
cells have microscopic
hairs that are moved by
outside forces such as
sound vibrations in the ear,
or water currents in the
neuromast organs found in
a fishs lateral line (right).
As the hairs move, their
cells produce nerve signals
that travel to the brain.

sensory input)
spinal cord



The spinal cord runs along a tunnel
formed by aligned holes in the
vertebrae, which protect it against
knocks and prevent kinks. Spinal
nerves, such as the sciatic nerve,
branch from it into the torso and
limbs. Cranial nerves branch directly
from the brain to the eyes, ears, and
other sense organs in the head, and
to the head and face muscles.

spinal column

In general, the hormone system works more slowly than the nervous
system. Hormones are chemicals made by groups of endocrine cells
scattered through various tissues or in separate endocrine glands. Each
type of hormonesome animals have more than 100spreads around
the body in the blood. It works as a chemical messenger to affect certain
parts known as its target organs or tissues, usually making them work
faster or release their products. Hormones maintain internal conditions
such as water balance and control
growth and development, the
reproductive cycle, molting or
shedding of body coverings, and in
some animals, metamorphosis
(drastic change in body shape).


Neurosecretions are made by nerve cells and
trigger actions such as hormone release or muscle
activity. A snails brain shows varying levels of
neurosecretion (yellow, red, and white) in response
to light levels sensed by the eyes, which makes the
snail active at night and restful by day.


The breeding behavior
of brown hares in spring
is triggered by rising levels
of reproductive hormones
from the sex glands
ovaries in females, testes in
males. Here, an unreceptive
female is boxing to fend
off a male.


Vertebrate nervous system

Vertebrate eyes are remarkably similar in form and
function. Light enters the eye through the central
pupil, the size of which is controlled by muscles
around the colored iris. The light is received and
processed by the retina at the back of the eyeball.


Being able to sense what is happening around them is vital
for animals survival. Among a myriad of other things, sensory
information helps them nd food and avoid predators; to know
when they are too hot or too cold, or are hurt; and to improve their
reproductive success by aiding the location and selection of mates.

What are senses?

A sense is the reception of a stimulus that is interpreted by the brain to
gain information about the external or internal environment. For example,
light is gathered by the eye and interpreted by the brain into an image.
The ve senses most familiar to humans are hearing, sight, taste, smell,
and touch. Most animals possess some or all of these senses to varying
degrees, but they may also have other senses, most notably echolocation
(the use of sound to locate objects), and electroreception and
magnetoreception (the abilities to detect electric and magnetic elds
respectively). In addition to being able to sense their surroundings,
animals require information about their own bodies, such as their
position and movement, which is sensed by special cells. For example,
thermoreceptors in the skin enable perception of temperature.





Jackrabbits have an acute sense
of hearing thanks to their huge
ears. Deep-sea squid have large
eyes that are sensitive to light
produced by bioluminescent
organisms. Butterflies have taste
receptors on their feet as well as
their mouthparts. Humpback
salmon have a keen sense of
smell that helps them find the
stream they hatched in. Lobsters
have long antennae and tiny hairs
all over their body that are
receptive to touch.


Senses and behavior

The senses play a vital role in animal behavior. Senses are required for
communication between animalsfor example, to hear and see sound and
visual signals, and to smell scent marks. They are needed to locate prey
for example, by sight, echolocation, or electroreception; to nd mates,
such as by homing in on pheromones or a mating call; and for navigation,
which may be mediated by sight, smell, or magnetoreception of the Earths
magnetic eld, or a combination of such cues. Animals may have evolved
particularly acute senses, or senses in specic ranges, according to the

Stellers sea eagle hunts by
sight. Its powerful binocular
vision allows it to spot prey and
accurately judge distance.

demands of their environment and behavior. Dogs have a well-developed

sense of smell, which they use to locate prey and communicate with one
another; bats can hunt at night in total darkness because of their ability to
use ultrasound for echolocation; many insects and birds can see ultraviolet
light, allowing them to detect
patterns on owers and plumage
that are invisible to the human eye;
and elephants can produce and
hear infrasound, permitting them
to communicate over many miles.
The brain integrates information
from all of the senses so that an
animal has a mental picture of
the world around it, enabling it to
react accordingly. For example, a
predator spotting possible prey will
Black-backed jackals greet one another by smelling
use sensory information to decide
scent glands located in the anal region. Smell is
important in identifying members of the pack.
whether or not to launch an attack.

Sensory systems
Each sensory system comprises sensory
receptors, neural pathways, and parts of
the brain involved in sensory perception.
Examples in vertebrates include the visual
system, where receptors of color and
brightness in the retina trigger nervous
spinal cord
impulses in the optic nerve, which travel to
the primary visual cortex for processing; and the
auditory system, whereby hair cells in the inner ear
receive vibrations caused by sound waves, triggering
nervous impulses in the auditory nerve, which travel
to the primary auditory cortex. The somatosensory
system detects pressure and touch; the gustatory
system senses taste; and the olfactory system receives
and processes information about smell. Each neuron
has a threshold, above which a stimulus will cause it
to re an impulse. The nature of the nervous impulses
triggered by the receptors provides the brain with
information about the location of
the stimulus, its intensity, and its
duration. For example, the longer
In vertebrates, such
as this seahorse, the
an object touches part of the body,
central nervous system
the more the receptor cells will
comprises the spinal
cord and the brain.
trigger the nerves.

senses 120

Touch and vibration

guard hair

Animals learn a great deal about their environment by using touch and sensing
vibration. They can feel for their food and communicate without the need for
sight or sound, skills that are especially helpful in the dark. The sense of touch
is facilitated by structures called mechanoreceptors that respond to stimulation.
Mechanoreceptors also provide feedback about an animals movement and
orientation so that it can adjust its position as necessary.
following its nose
As the star-nosed mole
hunts for soil-based or
aquatic prey, the 22
sensitive, fleshy tentacles
around its nostrils wiggle
in constant motion.

hairy legs
A spiders leg is covered
in hairs that respond to
touch and airborne
vibrations. The base of
each sensillum connects
with the dendrite of a
neuron, which transfers
a stimulus to the brain
for processing when
the hair is touched.

Touch receptors

mammalian skin

undercoat of
wool hairs

erector pili


Meissners corpuscles
Most arthropods detect touch and
in the upper dermis
vibration through sensory hairs
respond to light
touches, while the
called sensilla. These filaments
larger Pacinian
protrude through the exoskeleton
corpuscles, located
from where they are anchored in
around hair
deep in the dermis,
detect heavy, more
the epidermal layer below.
sustained pressure
Pacinian corpuscle
Movement of the hair triggers a
(pressure receptor)
and vibrations.
nerve impulse in an adjacent
receptor cell. Vertebrate hairs work in much the same
way: the base of the hair is located in a follicle with the
tip of a sensory neuron wrapped around the hair shaft
to receive and transmit the stimulus. In addition to hairs,
the epidermal (top) and dermal (lower) layers of the skin
contain a variety of different structures for detecting
touch, pressure, and vibration. Mechanoreceptors may
be concentrated to create a particularly sensitive area,
for example, on the nose of a star-nosed mole.

Motion detectors
Arthropod sensilla also detect
movements of air or water around
them, as do vertebrate whiskers
(vibrissae). These stiff, long hairs
are usually located around the nose
and mouth or above the eyes. They
are anchored in special follicles
called blood sinuses, which allow
even a tiny deflection of the whisker,
such as might be caused by a whisper of wind, to be
amplified and stimulate mechanoreceptor cells. Fishes
and larval amphibians have a different system to detect
movements in water. The lateral line is composed of
receptors called neuromasts, each of which has several
hair cells projecting into a gel-filled cap called a cupula.
The cupula bends in response to water movement, allowing
the animal to detect the direction of water currents.

lateral line
The lateral-line system is visible as a
faint line running down each side of a
fishs body. In sharks, rays, and many
bony fishes, the neuromast receptor
cells are located in a canal beneath
the skins surface. The canal connects
with the external environment through
a series of pores.

hair cell



multiple hairs
The hairs that
protrude above
a mammals skin
have a variety of
functions, including
sensitivity to
touch, insulation,
camouflage, and

Gravitational detectors
Many invertebrates possess structures called
statocysts that sense changes in orientation and
movement. A statocyst comprises a relatively heavy
ball, the statolith, inside a hollow sphere. The statolith
may be secreted by the statocyst or, as in lobsters,
it may be formed of sand grains collected from the
environment. The vertebrate inner ear also operates
like a statocyst. Otoliths, the equivalent of statoliths,
move against hair cells called cristae inside fluid-filled
ducts (ampullae) to detect gravity and acceleration.
scallops in motion
Scallops propel
themselves along by
expelling jets of water
from their shells. A
statocyst provides the
scallop with information
on its orientation,
prompting it to adjust
its path accordingly.
hollow cavity

Many species of small
bony fishes, such as the
white salemas (grunts)
seen here, form large,
dense schools as a
means of protection
against predators.
Their lateral-line system
enables the fish to move
in unison, and predators
find it hard to pick out an
individual target since the
school constantly moves
and makes sudden
changes in direction.

sensory hairs


bivalve statocyst
As the animal moves, gravity acts
on the statolith in the center of the
statocyst, causing it to stimulate the
sensory hairs of the receptor cells
against which it rests.


Vibration receptors are attuned

to vibrations that are transmitted
through a surface, usually the ground,
lower jaw
but also other surfaces such as tree
trunks, leaves, or spiders webs. The
detection of vibrations felt through the
environment is often a precursor to hearing the
vibrations of sound waves transmitted through the
air. In animals that lack a tympanic membrane (see
p.126), such as some amphibians and reptiles, sound
vibrations are transmitted through the body to the inner
ear. Animals that lack ears are able to feel sound by
detecting its vibrations, rather than hearing it. Many
mechanoreceptors, including insect sensilla and

signal passed
to inner ear

quadratic bone



bone conduction
Snakes receive groundborne vibrations by
pressing their lower
jaw to the ground. The
vibrations are conducted
to the stapes bone and
on to the inner ear.

Pacinian corpuscles, detect vibrations. Other vibration

sensors include lyriform organs on spiders legs, which
detect the vibrations of struggling prey trapped in their
webs, and Herbst corpuscles in the bills of wading
birds, which detect the vibrations of prey moving in the
sand. The ability to sense vibration is particularly useful
underground, where sound does not travel very far and
visual signals are of no use (see panel, right).

sensitive feet
This raft spider can
detect aquatic prey
by placing its front
two pairs of legs on
the surface of a
pool of water.
the cats whiskers
Whiskers are incredibly
sensitive to touch and air
movement. They permit
animals, such as this
leopard, to feel their way
and hunt in conditions
of poor visibility.

cAse sTudy

seismic signaling
Cape mole-rats communicate by drumming
their hind legs on the floor of their individual
burrows to create vibrations. During a
typical foot-thumping interaction, a male
and a female mole-rat will drum together
in synchrony. Their seismic signals travel
through the ground to neighboring burrows
more efficiently than sound.
foot drum



foot roll


foot-thumping session
When ready to breed, Cape mole-rats signal
their sex by drumming at different tempos in
order to locate a mate.


Taste and smell

Taste and smell are important senses for many animals, helping them
nd food and mates, communicate, and navigate. Both senses are
mediated by special cells called chemoreceptors, which bind with
specic chemicals then send a nervous impulse to the brain, where
the taste or smell is processed.

The sense of taste allows an animal to detect and identify molecules from
objects that come into contact with its gustatory (taste) receptors. These
sensory cells may be concentrated in different regions, for example, in the
mouth or on mouthparts, in the skin, or on the feet, depending on the animal.
They can be used to nd food and check that it is good to eat. For example,
ies have gustatory sensilla (hairs) on their feet with which they taste things
they land on. The sensilla are sensitive to different substances, such as
sugars, salts, or water. Taste is also used by some animals as a means
of chemical communication.

The whiskerlike barbels around the mouths and
noses of catfishes are covered with taste buds.
The barbels are used to locate food in the murky
waters at the bottoms of streams and rivers.

pore leading to
taste bud

sensory papilla


The surface of a mammals tongue (above)
is covered with sensory papillae, which
surround small pores that lead to barrelshaped taste buds beneath. Mammals detect
flavors with their tongues and some, such as
the brown hyena (left), will taste scent marks
to gain information about another individual.

The sense of smell is governed by olfactory receptors, which detect
odor molecules from objects at a distance. The molecules may be carried
on the air or in the water. On reaching an animal, the molecules bind to the
membrane of olfactory hairs (cilia). In arthropods, these are usually located
in pits within the exoskeleton or on bristly extensions of the exoskeleton.
In vertebrates, the cilia are usually located on the surface of the olfactory
tissue (epithelium) in the nasal cavity of the nose. Glands within the
epithelium secrete mucus that keeps the surface moist, helping trap
the odor molecules. Mammals, particularly rodents and carnivores,
have a good sense of smell. They use it to nd food, and many use
scent as a means of communicationto mark their territory, for example.

main olfactory bulb

olfactory epithelium
The main olfactory
epithelium is rich in
neurons that are receptive
to odor. They join to
form the olfactory nerve,
which connects to the
main olfactory bulb in the
forebrain. The vomeronasal
organ detects pheromones
and connects to the
accessory olfactory bulb.


Southern giant petrels have long tubular nostrils

with which they can locate food by day or night.

olfactory bulb

Eurasian water shrews have poor eyesight, but this is

compensated for by their acute sense of smell.

Emperor moth males can detect the scent of a

female 6 miles (11 km) away using their antennae.




Dogs are renowned for having a good sense of smell. Their sensitive noses have long
been employed by humans to assist in various tasks, such as tracking missing people and
detecting illegal substances. Recent research has shown that dogs can sense whether or
not a person has breast or lung cancer according to the smell of their breath. Cancerous
cells produce different metabolic waste products to normal cells, and it is these products
that the dogs are able to identify. The advantage of canine help is that cancers may be
identied and diagnosed earlier, leading to more effective treatment.

Many vertebrates have a Jacobsons or vomeronasal

organ, a patch of specialized olfactory epithelium, situated
in the roof of the mouth at the base of the nasal cavity.
This organ is particularly sensitive to airborne molecules
contained in scents such as pheromones, which are
used for chemical communication between
animals of the same species. Some mammals,
including most ungulates and felids, such as lions
and tigers, raise their heads and grimace when
testing the sexual receptivity of a female, a
behavior known as the Flehmen response.
Lizards and snakes use the Jacobsons organ to
detect chemicals such as those produced by
potential prey or a possible predator, transferring
the molecules from the air with their tongues.
This is why they constantly ick their tongues
when exploring their surroundings or when
they have been disturbed.


Tasting the air

Ungulates such as zebras (right) and
giraffes (below), show a characteristic
behavior when tasting the air for
pheromones. They pull the upper lip
back and draw air across the
vomeronasal organ. Through its use a
male is able to tell whether a female
is ready to mate.



extended tongue

retracted tongue

In snakes, the Jacobsons organ has two small
openings in the roof of the mouth into which the
tips of the snakes forked tongue are inserted
when it is withdrawn.

Snakes flick their tongues in and out to transfer
odor-bearing molecules from the air, water, or
ground to the Jacobsons organ. Elephants do
the same thing, but instead use the fleshy
finger at the tip of their trunks.



lens focuses
incoming light beam

Vision, or visual perception, is the way in which animals interpret information about
their environment using visible light. Some animals are only sensitive to the presence
or absence of light, while others can determine differences in the wavelength of
light, an ability known as color vision. The visual ability of an animal inuences
many aspects of its behavior, including feeding, defense, and courtship.

pigment cell

optic nerve
carries signals
to brain
light-sensitive cell
converts image
into a signal

What is an eye?

Compound eyes

Light is effectively parcels of energy called

photons, which travel in a wave, the frequency
(or wavelength) of which is proportional to the
energy they contain. An eye is an organ that
detects this light and translates it via nervous
impulses to the brain, where the information
is processed further. There is a wide variety
of eye designs across the animal kingdom,
relating to the different types of environments
that animals inhabit and the different behavioral
tasks they undertake.

African land snail eyes can only

distinguish between light and dark.

Most arthropods have eyes formed from multiple units

called ommatidia. Each of these consists of a lens to
focus light into a cell, a transparent crystalline cone to
funnel the light, and visual cells that absorb the light
and trigger a nervous impulse. The more ommatidia an
eye has, the more rened the resulting image. Each cell
is angled in a slightly different way, collecting light
from a slightly different area of the visual
scene. This makes compound eyes
excellent at detecting motion,
since the ommatidia are
consecutively turned on
and off as an object
passes across
the eld
of vision.

Cuttlesh have excellent eyesight

and distinctive w-shaped pupils.

Mandarin shes, like most tropical

shes, have good color vision.

Red-eyed treefrogs have a third

eyelid that helps keep the eye clean.

Chameleons can move their eyes

independently of each other.

Sourthern ground hornbill eyes have Tigers can see well at night because
long eyelashes that act as sunshades. of a mirrorlike layer in their eyes.


There are many benets to having two
eyes, both as a backup system should
one become damaged, and as a way to
increase the eld of view. But perhaps
the greatest benet is that they can be
used together to gather information
about distance. Eye position relates to an
animals behavioral need. Many animals
have eyes positioned on the sides of their
head, offering a very wide eld of view,
of up to 360, from which to watch for
predators. Most of this eld is monocular,
with little sense of depth. Predators,
however, typically have forward-facing
eyes, with a comparatively wide overlap
of the visual elds. This binocular vision
enables predators to judge distances
with great accuracy.

rear field of
binocular vision
blind spot
field of
forward field of
binocular vision


field of


field of

The multifaceted compound
eyes of the hovery are
composed of many thousands
of tightly packed ommatidia.
Different ommatidia have
different visual pigments,
which allowi the y to
see different colors of light.

crystalline cone
funnels light

Each ommatidium
sees a single
image, which the
animals brain puts
together, forming
a blurred mosaic.

Light enters a vertebrate eye through the transparent

cornea, which bends the rays. The light rays pass
through a uid-lled space to the iris, a pigmented,
muscular layer that can alter the diameter of its central
hole, the pupil, to control light entering the eye. The
light rays reach the crystalline lens, which can be
squeezed or relaxed to bend and focus the light rays
onto the retina at the rear of the eye, where an upsidedown image is formed. The retina is a layer of lightsensitive cells, which convert the information from the
light rays into nerve signals. The optic nerve carries the
signals to the brain, where an upright image is formed.


eye muscle moves

eye in socket

ciliary muscle alters

shape of lens

Vertebrate eyes

retina contains lightsensitive cells

light rays
from tree
enter eye

light rays cross

inside eye
crossed-over rays
produce an upsidedown image on retina
optic nerve

cornea bends
incoming light rays

The eyes of all vertebrates
share a common design, which
works like a camera, using a lens
to focus light and form an image.

lens changes
shape to focus
light on retina


There are two types of
photoreceptor in the retina.
Rods (stained yellow) are
sensitive in low light while cone
cells (stained blue) are involved
in color vision.

Color vision


Mantis shrimp have 12 different
spectral receptor types in their
compound eyes, sensitive to
wavelengths ranging from
ultraviolet to infrared.

Color vision is the ability of an animal to differentiate

between objects based upon the wavelength of light
they emit or reect. Cone cells contain pigments that
are tuned to absorb light from different regions of the
visible spectrum (see below), which the brain detects
and translates as color. The more cone pigments an
animal has, the more colors it can tell apart. Old World
monkeys and primates (including humans) have three
types of cone cell, while many shes and birds have
four. Most mammals have just two
cones and see in a similar way to a
color-blind human.
Cone pigments are classified by the wavelength of light they are most
sensitive to, for example short-wave (or blue light), medium-wave
(green light), or long-wave (red light).










In the animal
kingdom, color is
widely used for sexual
signaling. The red color
of a male mandrills
face conveys
information about his
age, rank, and
testosterone levels.

2 million

The number of different

surface colors that Old World Monkeys,
apes, and humans can see.

Beyond the visible spectrum

Humans are sensitive to wavelengths of
light in the range 400700 nanometers
(nm), but many animals, including birds,
insects, reptiles, and shes, can detect
short-wavelength light in the near-ultraviolet
(UV) range (320400nm). UV sensitivity is
used in many behaviors, including courtship
(see p.333) and predation (see p.228). An
unrelated ability is that of many animals, including
insects, birds, and cephalopods, to perceive polarized
To human eyes, this
light. Polarization occurs when, for example, sunlight is
flower is yellow, but
scattered by atmospheric particles to vibrate in a specic a UV-sensitive camera
reveals the patterns
plane. The way in which sunlight is polarized is related
that guide insects to
the nectar at its center.
to the suns position, so animals can use it to navigate.

Noctuid moth 1,000240,000


There is a vast array of hearing ranges
across the animal kingdom. Humans
can only hear sounds with a frequency
above 20 hertz and below 20,000
hertz (this sound range is called sonic).

Dolphin 110150,000



The ability to hear allows animals to sense the rustles

of approaching prey or respond to a mating call. In order
to hear, an animal must have a mechanism for detecting
the vibration of sound waves; sound receptor cells have
tiny hairs that are deected by sound waves, triggering
nervous impulses that are processed in the brain.

ULTRASONIC Above 20,OOO hertz (20kHz)

SONIC 2020,000 hertz


INFRASONIC Below 20 hertz (Hz)

Sound is produced by pressure waves that create disturbance in a

medium (usually air or water) detectable by hearing apparatus. Pressure
waves that are transmitted through a solid medium, such as the ground,
are usually referred to as vibrations. The number of oscillations
or vibrations per second is called the frequency of the
sound and is measured in hertz (Hz). The frequency
of the sound wave determines its pitch: a highpitched sound, such as a bird whistling, makes
the air molecules vibrate backward and forward
more times each second than a low-pitched
sound, such as the croak of a bullfrog.

Sounds emitted and perceived by animals can
be broadly separated according to frequency
into infrasonic (very low frequency), sonic,
and ultrasonic (very high frequency).

Bat 10100,000



For each species of
frog, the size of the
tympanic membrane
(behind the eye), and
the sensitivity of the
females ear, is related
to the frequency of
the males call.


ULTRASONIC Above 20,OOO hertz
SONIC 2020,000 hertz

The antenna of the
male mosquito is most
sensitive to sound around
380 hertz, the frequency
produced by a female
mosquito in flight.
Movement of the antenna
in response to a sound
wave stimulates the
auditory sense organ
at the antennas base.

Cat 7064,000

INFRASONIC Below 20 hertz

Dog 2045,000






Cricket 9511,000
Elephant 120,000
Human 2020,000


Sound detection
Movement-sensitive sensilla (hairs)
in insects detect the vibrations
of sound waves. Some insects,
including grasshoppers and some
moth species, have tympanic ears, which are more similar
in structure to those of frogs, some reptiles, birds, and
mammals (although grasshoppers have eardrums in their
abdomen rather than their head). Tympanic ears have a
thin membrane that transmits sound vibrations. In many
frogs, the membrane is visible on the sides of the head,
while in other animals it is obscured by elaborate ear
structures that improve sound reception. Some sh use
their swim bladder as a hydrophone to transmit sound
waves through small bones
that connect it to the inner
A typical mammalian ear comprises
ear. To communicate over
the external ear (pinna or auricle),
long distances, elephants
which leads to the tympanic
make infrasonic rumbles that membrane; the middle ear, an airthey can detect through their filled cavity across which sound is
transferred by the auditory ossicles;
feet and trunks as well as
and the inner ear, where the sound
receptor cells are located.
with their ears (see p.467).


ear pinna





Sound localization
The presence of two ears allows an animal to determine the origin of a
sound through localization. Unless the animal is facing directly toward
the sound source there is a delay between the sound arriving at the ear
closest to and farthest from the source. The brain uses this split-second
delay to calculate the direction from which the sound came. To gain
information about the height of a sound, the two ears must be
asymmetrically placed, as in the barn owl.
Its heart-shaped facial disk of feathers
acts like a radar dish, collecting sound
and guiding it to the eardrums
within the skull.
In some species of owls, the right ear cavity is
usually higher (and larger) than the left, so a
sound coming from below the owls line of sight
will arrive sooner at the left ear.

internal cavity
of right ear

internal cavity
of left ear

A barn owl localizes sound so well that it
can hunt in total darkness by listening for the
movement of its prey, usually small rodents,
rustling through the undergrowth.
A barn owl can detect
a time lag of 30 msec
between sound
reaching each of its
ears. This information
is processed by
10,000 spacespecific neurons.
To pinpoint its prey,
the owl moves its
head until the sound
reaches both ears
as the same time.


movement of
mouse generates


30 msec
The huge ears of
the bat-eared fox
gather and amplify
the sounds of its
insect prey moving
beneath the surface
of the soil.

The loudest animal

on the planet is the
blue whale, with
a call of up to 188
decibels. A jet
engine reaches
only 140 decibels.

Wrens are tiny, but their songs can

be heard from hundreds of yards.

Rattlesnakes shake the modied

scales at the tip of their tails.
cartilage ring
clavicular air sac

The syrinx is located where
the trachea branches into
the two bronchi. Air passing
over the pessulus and
the external and internal
tympaniform membranes
causes them to vibrate,
producing sound.
external tympaniform

bronchi lead downward
to lungs

Sound generation
Animals produce sounds in many different ways. Perhaps the
simplest form of sound production is hitting something, often
the ground, with a body part. For example, ants stamp their
feet, ddler crabs bang their claws, and mole-rats thump with
their heads or feet. Many insects and some other arthropods
generate sound by rubbing parts of their body together, a
method called stridulation (see p.457). Terrestrial vertebrates
make vocalizations by using the movement of air in the
respiratory system. For example, vibrations of the vocal
cords of the larynx produce a wolfs howl, while bird calls
and songs are produced by the syrinx (see left).

Wolves howl to signal ownership of

their territory to other wolf packs.

Echolocation, also called biosonar, is the ability to locate objects through the use of
sound. Animals that echolocate form an image of their surroundings from returning
echoes, allowing them to pick a path through vegetation or a cave system, or to home
in on their prey in the dark. The majority of bats use very high-frequency (ultrasonic)
calls to echolocate, whereas dolphins and most other echolocators use clicks.

Echolocation calls


Sonograms depict the frequency range, duration,
repetition rate, and shape of the echolocation call.
Calls from common pipistrelle, Daubentons, and
Leislers bats differ in all of these attributes.


Echolocation calls can be sonic,

that is within the range of human
hearing, but most echolocators use
very high-frequency, or ultrasonic,
Leislers bat
Daubentons bat
calls, which are measured in
kilohertz (kHz) . The types of calls
are incredibly diverse. Different bat
species produce different calls
according to the habitats in which
0 20
100 140 180 220
they forage. A fast-ying species
that hunts in the open air uses
relatively low-frequency calls that travel farther ahead of the bat, whereas
a slower-ying species feeding in a cluttered habitat, such as among
trees, requires detailed resolution gained from higher-frequency calls.
Bat echolocation calls typically range from 20 to 100kHz, whereas some
species of toothed whales use even higher-frequency clicks, with those
of bottlenose dolphins reaching
220kHz. Sperm whale clicks range
from 40Hz to 15kHz.

The majority of bat species are sophisticated

echolocators, producing their ultrasonic calls in
the larynx. Most of these bats emit the call through
the mouth, but some send it out through their nose,
which has lead to the evolution of elaborate nose
ornamentation. Many bats have large, forwardfacing ears with a projecting lobe called the tragus
that helps receive returning echoes. Bats usually
emit one call for each wing beat when ying. Their
calls are so loud that bats risk deafening themselves.
To avoid this, muscles in the middle ear contract
when the bat calls and then relax to allow the
returning echo to be heard before the bat calls again.
When potential prey has been located, the rate of
calling increases as the bat homes in on its target.
This bat is named for its small
size and horseshoe-shaped
nose-leaf. Furrows in the noseleaf help shape and focus the
beam of sound as it is emitted
through the nose.


This sonogram of a single sperm whale
echolocating shows it emitting a steady series of
clicks while searching for prey. Sperm whales eat
mainly octopuses and squid, even giant squid.

Up to 65 ft (20 m) in length, the sperm whale is the
largest toothed whale. Its huge, square-ended head
contains a spermaceti organ, a large mass of waxy oil,
which focuses echolocation clicks into a beam.





This species belongs to a
genus of Old World fruit bats
that echolocate by producing
pairs of sharp clicks with their
tongue. They have good eyesight
and use this relatively crude form
of echolocation to find their way
in dark caves.



Toothed whales


Dolphins produce rapid bursts of ultrasonic clicks
to locate prey. They also make an array of other,
lower-frequency sounds to communicate with each
other, including whistles and squeaks.

Echolocation allows toothed whales to pursue agile

prey, even in dark or turbid waters. Porpoises and
dolphins produce ultrasonic clicks by forcing air
between phonic lips in their nasal passages. The lips
open and close causing the surrounding tissue to
vibrate and form sound waves. These bounce off the
bony cribriform plate at the front of the skull and are
focused into a beam by the melon, which the hunter
aims at its prey. Sperm whales have a different
anatomy for producing echolocation clicks. The left
nasal passage is used for breathing, while the right is
used for sound production. Sound waves pass through
an oil-lled spermaceti organ, rebound off an air sac at
the rear of the head, and are focused into a beam of
sound through several fatty lenses. Experiments with
bottlenose dolphins have shown them to be extremely
sophisticated echolocators, capable of identifying
submerged objects by size, shape, or composition,
for example. This enables them to learn the echo
signatures of their preferred prey species.

vestibular air sac

phonic lips
melon pointing at fish

cribriform plate

nasal sac
inner ear
acoustic window

fat-covered, oil-filled sinuses in

lower jaw
mesorostral cartilage

Only two families of birds are known to echolocate:

the cave swiftlets of Southeast Asia and Australia and the
oilbirds of South America. Both roost deep inside caves,
where the ability to see is not of much use, so instead
they use sound to avoid bumping into the rock walls
and each other, and to nd their nests. They produce
low-frequency clicks of less than 15 kilohertz (kHz), by
contracting muscles around the vocal organ, or syrinx
(see p.127), which cause the tympaniform membranes
to vibrate. Their echolocation is low resolution and both
groups feed by sight; swiftlets on insects during the
day and oilbirds on oil palm and laurel fruits at night.










The swiftlets clicks are emitted in pairs around 18

milliseconds apart. The first click has a frequency
of most energy between 3.3 and 5.5kHz and the second,
louder, click has a frequency of between 4.1 and 5.5kHz.


As the Australian swiftlet flies deeper into the
darkness of its cave, it lessens the interval between
the pairs of clicks it emits to gain more information
from faster-returning echoes.

False vampire bats such as the ghost bat of northern Australia
have pointed leaf-shaped nose-leaves through which they
direct their calls at both invertebrate and vertebrate prey. Their
large ears meet in the middle and each has a prominent tragus.

Other mammals

sound wave emitted

by the dolphin
echo returning to the dolphin
from the fishs swim bladder

swim bladder
fish swimming
ahead of dolphin

Dolphins use the melon, an oily lump of tissue behind the
forehead, to focus their outgoing sound waves into a beam.
Echoes returning from a fish in the dolphins path are transferred
through oil-filled sinuses in the lower jaw to the inner ear.

The streaked tenrec searches for worms and grubs
in the Madagascan rain forest floor, making a series of
low-frequency tongue clicks to locate its prey.

Bats and toothed whales have by far

the most sophisticated echolocation
among mammals, although some
nocturnal insectivores also use primitive
echolocation to explore their habitats
and catch their insect prey. Shrews
produce ultrasonic calls from the larynx
whereas the tenrecs of Madagascar
use less specialized tongue clicks. In
addition, both may communicate with
other individuals at ultrasonic frequencies.
The aye-aye, also from Madagascar, uses
sound to locate its prey too, but in a very
different manner. It taps a tree trunk with
its long, bony middle nger and listens
for the larvae of wood-boring beetles
moving underneath (see p.247).




Electricity and magnetism

Many animals possess the ability to sense electric and magnetic elds. Some detect the
electrical signals generated by the muscles of other animals, while others produce their
own electricity, either to use in prey location, navigation, or communication, or as a
means of defense or prey capture. Animals with a magnetic sense use the Earths
magnetic eld for navigation or orientation.


The duck-billed platypus has about 40,000
electroreceptors in its bill, which sense
the electric fields of prey concealed in
muddy river bottoms.

The short-beaked echidna inhabits
drier habitats than the long-beaked
echidna and has only around 400
electroreceptors at the tip of its
beak. It uses them to locate prey
when conditions are wetfor
example, when it rains.


Animals that are capable of detecting electrical

impulses but do not generate them are said to show
passive electroreception. Those that both receive
and generate electrical signals are capable of active
electroreception. Electroreception is more common
among animals living in water than on land because
of the capacity of water to conduct electricity. The
majority of electroreceptive animals are shes, but
monotreme mammals also possess the ability. For
example, the long-beaked echidna, which
lives in wet tropical forest, has around
2,000 electroreceptors in its beak,
which it uses to track down
earthworms and other soildwelling prey. This use of electric
elds to nd food is known as
electrolocation. The structures
responsible for electroreception
are called ampullary receptors. The
electroreceptive sense organs found
in sharks and rays are known as ampullae of
Lorenzini after the Italian anatomist Stefano Lorenzini,
who described their structure in 1678; their function
remained a mystery until the 20th century.

Weakly electric fishes,
like this elephantnose
fish, generate small
electrical pulses of less
than 1 volt. They then
detect distortions in
the electrical fields
they produce and use
these for navigation,
location of prey, and

pore in

of Lorenzini


sensory cell at
base of canal


An ampulla of Lorenzini consists of a pore in the skin
opening to a gel-filled canal. It works by detecting the
difference in voltage between the pore and the base of
the canal, allowing the shark to sense electric fields such
as those generated by the muscles of a fish.

Electrogenesis is the production of an electrical discharge by an animal.
Strongly electric shes, such as the electric eel, electric catshes,
and electric rays, produce much higher voltage pulses than
weakly electric shes. The organs of electrogenesis, called
electrocytes or electroplaques, are usually modied muscle cells or
occasionally nerve cells. The electric eel has between 5,000 and 6,000
electroplaques stacked in series in its abdomen, enabling it to generate
huge shocks of up to 600 volts to stun prey when hunting. The electric
eel also uses lower voltage pulses of around 10 volts for navigating and
detecting prey. Electric rays use the electric organs in their heads to
electrocute prey or stun a potential predator. The size of the electric
discharge varies from 8 to 200 volts, depending on the species. The
electric catsh produces its electrical discharge in its skin.


nerve controlling
electric organ

electric organ

Despite its name and appearance, the
electric eel is not a true eel but a type
of freshwater fish known as a knifefish.
The electric ray has a pair of electric organs on
either side of its head, which can produce an
electrical discharge of up to 200 volts.

Sharks snouts are covered with electroreceptors
called ampullae of Lorenzini, which appear as dark
spots. Sharks may also orient themselves relative to
the Earths magnetic field by using their own electric
field and those generated by ocean currents.

Sharks can detect tiny

electrical signals as weak
as 15 billionths of a volt

Some animals are able to perceive
the Earths magnetic eld and use it
for navigation. This ability to detect
changes in the magnetic eld is called
magnetoreception. At least two different
mechanisms may be involved. The rst
involves crystals of magnetite, a magnetic
form of iron oxide, which has been found in
many speciesfor example in the upper
This scanning electron micrograph
mandible of pigeons, in the abdomen of
of a particle of magnetite reveals the
honey bees, and in the head of trout.
minerals classic octahedral shape.
It is the most magnetic of all the
Clusters of the mineral are thought to be
naturally occurring minerals on Earth.
sensitive to changes in magnetic intensity.
The second mechanism is known as the
radical pair model. It is thought that a magnetic eld alters the spin of
electrons in a specialized photopigment called cryptochrome, allowing the
direction of the eld to be determined. It is possible that birds might use
both mechanisms simultaneously: magnetite in the bill to sense the intensity
of the magnetic eld and so locate magnetic north; and cryptochrome in
the right eye to sense the direction in which they are ying.


Experimental alteration
of the magnetic field
around the lofts of
homing pigeons has
been shown to disrupt
their homing ability,
lending weight to the
theory that they
use magnetic fields
when navigating
their way home.


Magnetic termite
mounds are aligned
northsouth so that
their broad, flat sides
face east and west,
which helps keep
the mound at
86 F (30 C).
Experiments have
shown that
termites use the
Earths magnetic
field to orient
their mounds.




Home ranges
and territories




Animal architects

The Antarctic ice floes are among the most extreme
habitats on the planet. Temperatures regularly
plunge below freezing, yet chinstrap penguins make
their home in this spectacular landscape with access
to plenty of food in the sea and few predators.


An animals living space should contain all an animal needs to survive.
The living space might be very small if the animal is xed in position, such
as an adult barnacle, or very broad if the animal moves around a lot in the
course of feeding or if it is a species that migrates.

Habitat choice
Put simply, an animals habitat is its living space. Animals choose to live in
a habitat that enables them to maximize their tness by providing all they
need to produce offspring and maximize their own survival. Animals show
adaptations, both physical and behavioral, to their preferred habitats.
For example, coal tits prefer to live
in pine forests while blue tits like
oak woods. Young coal tits are
better adapted to foraging among
pine needles, whereas blue tits are
more adept at searching for food
among oak leaves.

The Arctic fox (left) has thick white fur that
provides camouflage and insulation against
the cold. By contrast, the fennec fox (above)
has sandy brown fur and large ears that
dissipate heat to keep it cool.


An animals home range contains places for it to sleep, breed, and feed.
It may spend the majority of its time in one or more core areas of the range
and venture to the periphery only occasionally. Animals may disperse from
their home range into another or migrate periodically between different home
ranges, which can be separated by vast distances.
Many animals defend all or part of their
home range as a territory, either year


Arctic terns maintain breeding territories in Greenland
during the northern summer. A colonys home range
extends 1.9 miles (3 km) out to sea, where the birds
feed on small fishes. In fall, the terns migrate south
to the edge of the Antarctic ice sheet, where they
overwinter, moving south with the ice as it melts.





Home range









Migration route

round or just at certain times. For example, they may guard a mating site
during the breeding season, a particular seasonal food source, or a shelter
such as den or nest. Territories are often marked to warn neighboring
animals of the territory owners presence. These behaviorsdispersal,
migration, and territorialityhave costs in terms of predation risk, energy,
and time. In order to be worthwhile, the benets must outweigh the costs.

Surviving extremes
Around the world, animals cope with an astounding variety of environmental
conditions, and they have evolved a wide range of physiological and
behavioral adaptations to enable them to survive. Some animals possess a
natural antifreeze and can survive being frozen; others have thick insulating
blubber, fur, or feathers to avoid becoming chilled. Many animals hibernate
during cold periods or estivate during warm ones by reducing their
metabolic, heart, and breathing rates to the bare minimum until conditions
change and they can be active again. And some species can survive on
very little water by storing it or obtaining it only from their food.


Home range, territory, and migration


An Arctic tern is very protective of its
nest and chicks. It will fly at a predator,
such as a herring gull, that ventures
near its territory, attacking the
imposter with its sharp bill.

Emperor penguins live in Antarctica, where
temperatures reach -40 F (-40 C). Arctic
beetles also withstand -40 F in the wild, but have
been chilled to -125 F (-87 C) in a laboratory.
Pompeii worms live around deep-sea
hydrothermal vents in the Pacific Ocean, where
water is heated to 570 F (300 C). The worms
hold their heads in cooler water than their tails.
Peringueys desert adder lives in the Namib
Desert in Namibia, which receives less than 3 8 in
(1 cm) of rain each year. The snake obtains most
of its water requirements from the lizards it eats.
Yaks roam up to 19,700 ft (6,000 m) and
red-billed choughs up to 26,000 ft (8,000 m)
above sea level in the Himalayas. Vultures have
collided with planes at 36,000 ft (11,000 m).
The fangtooth fish is found at depths of 16,400 ft
(5,000 m). In addition to high pressures, deep-sea
fishes must cope with cold water temperatures,
reduced oxygen, and little to no ambient light.
Brine shrimps live in hypersaline lakes where
salinity may be 25 percent (normal seawater
is around 3.5 percent). The shrimps and algal
blooms color the water red and green.


Home ranges and territories

A home range is the place where an animal lives. Whether an animal defends all or part
of its home range as a territory or coexists peacefully with others on an undefended
home range depends on whether the benets of being territorial outweigh the costs
of territory defense. Some species become territorial for part of the annual cycle and
are nonterritorial occupants of a home range for the rest of the year.

Home ranges


A home range contains all areas used by an animal

during its everyday movements. It should be large
enough to contain food for the animal and its young,
and other resources such as shelter and water. Home
ranges consequently vary in size according to resource
availability. Polar bears, for example, have huge ranges
of up to 48,250 square miles (125,000 square km)
because their prey is sparsely distributed. An animals
home range might overlap with those of others, or
it may travel beyond its own home range to nd a
mate. Home ranges can be three-dimensional;
plankton and sh move up and down within
the water column, while bats and
vultures soar into the sky.


In 1958, Dutch ethologist Niko Tinbergen showed that digger wasps
nd their nest entrance by remembering the conguration of objects
around it. After emerging from her burrow, the female wasp ies
around the landmarks, in this case a circle of pinecones. If the circle
is moved once the wasp has left to hunt she will search for her nest
in the location indicated by the cones when she returns.
wasp learns
location of

have been


Many animals defend territories against intruders. Territorial boundaries are
often marked to announce the owners presence, thus avoiding potentially
harmful confrontations between neighbors. Marking may involve visual
displays, scent, or audible signals. An incursion into another animals
territory can escalate to combat
if the intruder persists. In social
species, larger groups often win
territorial disputes. The advantage
of territoriality is that the territory
holder can monopolize resources
such as food or a good breeding
site within the territory.
A northern gannets territory comprises its nest and
a small space around it. It viciously attacks any bird
other than its partner that ventures within reach.

Dispersal occurs when an animal leaves one home range or territory to
establish its own elsewhere. It may be forced to depart because changing
environmental conditions have rendered its current habitat unsuitable.
Alternatively, an animal may leave an area to avoid competition with rivals
for food or other resources. Dispersal is also a mechanism whereby
animals avoid breeding with close relatives. In many species, the males
disperse while the females stay at or close to their birth territory.
In some cases, once
young animals become
independent, they are
chased away by their
parents. These behaviors
ensure that the species
does not become inbred.
Adult black bears rub their backs, shoulders, and
the backs of their heads on trees to scent-mark
their territorial boundaries and to advertize their
presence to prospective mates.

Newly hatched garden spiders disperse
in different directions by walking or
using a strand of silk to catch the wind.

Keeping a distance
Dancing white lady spider

The common limpet grinds circular scars

with the edge of its shell into the rocks
on which it lives. When covered with
seawater at high tide, it moves away
from its scar to forage on algae. It returns
before the water recedes by following
chemical cues in a mucus trail it leaves
behind. Settling in the scar helps the
limpet create a strong seal to avoid
both predation and dehydration when left
exposed on the rocks at low tide. Limpets
play an important role on rocky shores by
keeping the substrate clear of algae,
allowing other organisms to colonize.

The dancing white lady spider inhabits

a burrow protected by a trapdoor in the
sands of the Namib Desert. Each individual
has a territory with a radius of 314 934 ft
(13 m), depending on the sex and age
of the spider. These spiders will apparently
resort to cannibalism in order to defend
their territories. It is hardly surprising then
that a male spider walking through another
males territory while out searching for a
mate will drum the sand to announce his

arrival, using all eight of his legs and even

his body. A male may travel up to 300 ft
(90 m) across the dunes during a nightly
excursion, crossing the territories of
between one and ve other males.
However, it may encounter only one or
two females on its journey. Immature and
female spiders tend to stay put in their
foraging territories.

10 ft

maximum radius of
a male dancing white
lady spiders territory
around its burrow.
Male dancing white lady spiders drum their feet
on the sand when they are close to other males,
causing rivals to withdraw or keep out of the way
of the signaling spider.

Leucorchestris arenicola

a 1 2 11 2 in (1.53.5 cm)
b Namib Desert, Namibia.
c Creamy white spider.

Patella vulgata

a Up to 214 in (6 cm) (shell diameter)

b Temperate rocky shores throughout Europe, from

Norway southward to Portugal.

c Conical, gray-white shell, sometimes tinged with

yellow or covered in algae. Muscular foot underneath
contracts and prevents limpet from being lifted.

Perch patrol
Common whitetail
Mature male common whitetail dragonies
develop a whitish bloom known as
pruinescence on their blue abdomens. In
territorial displays, dominant males raise
their abdomens to intruders as a threat,
while less dominant males lower theirs as
a sign of submission. Common whitetails
guard perches from where they patrol their
territory, a 3398 ft (1030 m) stretch of
waters edge, for several hours a day. Males
with larger territories have greater success
in mating with females that visit the water
to breed. They also defend their territories
against other species such as the cardinal
meadowhawk, as pictured below.
Libellula lydia

a Up to 2 in (5 cm)
b Wetland habitats throughout North America.
c Males have a light blue abdomen, females a brown

abdomen. Dark brown band down center of wings.

Fighting for position

Poplar petiole gall aphid
In spring, female poplar petiole gall aphids
hatch from eggs that have overwintered on
cottonwood trees. Each aphid selects a
leaf and starts feeding at the base, causing
a hollow ball of tissue called a gall to form.
The female moves into the gall, where she
gives birth to a brood of winged females,
the result of asexual reproduction.
These females disperse and continue to
reproduce. The last generation of the year
contains both male and female aphids,
which reproduce sexually and lay eggs that
overwinter. The females are extremely
territorial and may spend several hours, or
even days, ghting with one another over a
leaf. Large leaves are preferred, presumably
because they contain more sap on which
to feed. Defeated and small females must
accept smaller leaves or less favorable
positions closer to the center of large leaves.
Where three females occupy the same leaf, the
reproductive success of the female closest to the
base is always greater than that of the female farthest
away. On average, the female at the base produces 138
young, the middle female 75, and the most distant female 29.


Aphid galls are typically
located at the base of
the leaf where it joins the
petiole (stalk). Each gall
contains new aphids,
which are produced
parthenogenetically by
the female that created
the gall. They eventually
emerge to establish their
own galls elsewhere.

female farthest
from base has
least young
gall of
female at base
most young

Pemphigus populitransversus

a 116 1 8 in (1.53 mm)

b Eastern cottonwood trees in Illinois,

Missouri, and Utah, US.

c Green insect with a pear-shaped

abdomen. There are both winged and
wingless forms.


Home scar
Common limpet

ritual combat
Malaysian giant ant majors engage in ritual
fights at tournament sites on their territorial
boundaries. Two ants stand on their hind legs
and box one another with their forelegs. The
ant that holds its legs up longest and throws
its opponent off balance wins the bout.

Watery crib
Harlequin poison frog

Fighting colors
Common side-blotched lizard

The female harlequin poison frog gives

each of her tadpoles its own pool in the
base of an aerial plant. Water collects
where the leaves join, and the female
drops one tadpole into each reservoir. The
tadpoles are kept apart because they tend
to be cannibalistic. The adult frog returns
every other day to deposit an unfertilized
egg into the pool as food for her developing
offspring. The young frog emerges from its
aquatic nursery after three months.

Male common side-blotched lizards have three different strategies

when it comes to mating. Orange-throated males are territorial
and aggressive. They are dominant over both blue- and yellowthroated males, and consequently mate with the most females.
Blue-throated males are also territorial and guard their mates
carefully. They are able to chase yellow-throated males away, but
often lose out to orange-throated males unless they cooperate to
protect their mates. Yellow-throated males do not defend territories
at all; instead, they attempt to
Uta stansburiana
sneak past the territorial males
a 11 2 21 2 in (46.5 cm)
to mate. Early in the breeding
b Desert and semiarid areas of Pacific
season, before the males
North America and north-central Mexico.
coloration has developed,
c Females are patterned brown and
competitions for the best
white; males are mainly brown/gray but
have three color forms (orange, blue, and
territories are decided on the
yellow) during the breeding season.
basis of size rather than color.

casE sTudY

rocks count
Researchers tested the theory that male common side-blotched lizards
in a high-quality habitat maintain a smaller territory than those in a
poor habitat. They altered territories so that some males had valuable,
sun-warmed rocks taken away and others were given more. Males in
areas with fewer rocks expanded their territories to compensate for
the poorer-quality resource, while those with more rocks contracted
the size of their territory, but still managed to attract a higher density
of females.
extra rocks
provide more
basking sites

beFore alteration

Dendrobates histrionicus

a 11 2 in (2.54 cm)
b Tropical rain forest of western Ecuador and parts
of Colombia.

c Many color forms exist, including black markings

over a base of yellow, orange, white, red, or blue.

rockS added

Soaking up the Sun

Male (right) and female (left) common
side-blotched lizards bask on a rock
that has been warmed by the sun to
absorb the heat contained in the rock.

Living space 140

border guards
Malaysian giant ant
territorial behavior is well developed in the
Malaysian giant ant. colonies defend their
borders against ants of the same species
and also enter into violent combat with
ants of different species. colonies of
Malaysian giant ants may contain around
7,000 workers divided among 8 to 14
underground nests. A colonys territory
extends up into the tree canopy and may
cover an area of 86,100 square feet (8,000
square meters). Barrack nests near the
territory borders contain a high proportion
of major workerslarge soldier ants that
take part in ritual combat with other
Malaysian giant ants during boundary
disputes. these fights can take place
over a number of days, or even weeks,
with majors squaring up to one another
repeatedly each night. the majors also
patrol trunk trails through the colonys
territory, attacking impostors of their own
and different species, and serve
as sentinels at points known
as bridgeheads. these are
usually at the base of
tree trunks that give
access to the canopy.




rain forest habitat

Malaysian giant ants
build subterranean nests
in the forest floor. A few
ants are active on the
ground during the day,
but hordes of workers
travel into the canopy at
night to forage.
Camponotus gigas

a 34 in (2 cm) (worker); 114 in (3 cm) (soldier)

b Rain forest from lowland peat swamps and

mangroves to mountain forests of Southeast Asia, from

Sumatra north to Thailand.

c Large ant with a black head and thorax and a

brown rear abdomen.

Camponotus gigas nests

Camponotus festinus nests
Other Camponotus species nests
territorial boundaries
Distinct territorial boundaries are
maintained between different
Malaysian giant ant colonies and
between them and colonies of
other large, nocturnal ants
with which they compete.

Oecophylla smaragdina nests

Camponotus gigas sentries
Camponotus festinus sentries

aggressive defense
Titan triggerfish
the titan triggerfish is a large, rhomboid-shaped tropical reef fish
that defends its territory aggressively during the breeding season.
if approached, it repeatedly
swims at the intruder until
the intruder retreats. When
threatened while feeding on
the reef, a triggerfish may hide
between rocks and erect the
spines on its back, bracing itself
in position and thus rendering it
immovable. As the largest of the
triggerfishes, the titan is usually
dominant to other species in
competitive encounters.
Balistoides viridescens

a Up to 30 in (75 cm)
b Coral reefs of the Indo-Pacific Ocean.
c Heavily scaled fish. Main body

coloration can include yellow, black,

green, and dark gray.

d 400
nest site
Titan triggerfish typically build their
nests in sandy patches of coral rubble
on the seabed. They use jets of water
to excavate a chamber in the sand in
which the female lays her eggs.


nasty bite
The titan triggerfish is notorious for
attacking divers and snorkelers that enter
its territory while it is guarding eggs. It will
attempt to escort people away, but it has
been known to knock divers unconscious
and inflict bites with its strong teeth. The
bites can be poisonous if high levels of the
ciguatera toxin are present (this accumulates
in the fish as a result of eating contaminated
prey, including shellfish). The fishs territory
encompasses a cone-shaped body of water
above the nest, so to retreat safely, intruders
should swim away to the side, not upward.

nesting platform
Garibaldi damselfish
Male garibaldi damselfishes trim algae into a circular nesting
platform, to which they entice females by making a loud clicking
noise. Females are very selective and tend to choose nests with
a thicker layer of algae and that already contain eggs at an early
stage of development. this appears to be because male
garibaldis often cannibalize a
single clutch of eggs and they
Hypsypops rubicundus
will guard a large clutch more
a Up to 12 in (30 cm)
aggressively than a small one.
b Coastal reefs and kelp forests from
they vigorously defend the
central California to southern Baja
California, Mexico.
eggs for the two- to threeweek period between laying
c Adults are bright orange with a deeply
forked tail. Juveniles have blue spots.
and hatching.


Canyon soaring
Andean condor
The Andean condor soars at altitudes of
up to 18,000 ft (5,500 m) while scanning
the ground for carcasses to scavenge. It
has a large home range and may travel up
to 125 miles (200 km) in a day. This birds
excellent eyesight enables it to spot food
from several miles away, and it often uses
the presence of other vultures to reveal the
location of a potential meal, perhaps a
dead farm animal, wild deer carcass, or
even a beached whale. Condors rarely ap
their wings when ying; instead, they ride
on updrafts of warm air, circling with their
wing tips bent upward. Andean condors
nest at 9,85016,400 ft (3,0005,000 m)
above sea level on rocky ledges. Like their
relative, the California condor, they have
in the past been subjected to persecution,
which has caused their numbers to fall.
Captive breeding and raising programs
aim to improve their breeding success
to secure the birds future.
These Andean condors are soaring on
an updraft in the Colca Canyon in Peru.
They typically hunt in the early morning
and late afternoon.

Pecking distance
Black-browed albatross
Like many seabirds, black-browed
albatrosses are colonial, coming together
in huge nesting colonies during the
breeding season. They are forced to nest
together because of the limited availability
of islands suitable for breeding in the
Southern Ocean, and also because
communal living decreases the risk of
predation of eggs and chicks. The optimal
nesting density leaves them just out of
pecking reach of the bird next door. The
colony therefore appears to be evenly
spaced across the tussock-grass plateau,
each bird the ruler of its own small territory.
When away from their nests, black-browed
albatrosses range over hundreds of miles
of ocean, feeding on shes, squid, and
octopuses. This species is ofcially listed
as endangered because of the high
number of deaths caused by long-line
shing (see p.155).

Immature birds
may spend up to
seven years at sea
before returning to
establish a territory
of their own.
Thalassarche melanophrys

a 3137 in (8095 cm)

b Islands and open waters of the Southern Ocean.
c Mainly white bird with dark gray upper wings. Yellow

to dark orange bill and dark eyestripe above each eye.

d 253, 403
The worlds largest breeding colony of black-browed
albatrosses is on Steeple Jason Island in the
Falklands. A pair raises just one chick each year, so
their population is slow to recover from decline.

white ruff of

Andean condors
become sexually mature at
four or five years of age. Females
lay one or two eggs in alternate years.
Vultur gryphus

a 4414 ft (1.21.3 m)
b Open grassland and rocky areas of the Andes and

Pacific coast of South America.

c Large black vulture with featherless black and

red head and neck. Males have large white patch
on their wings.

The Andean
condor has the
largest wing area
of any bird.

The New Holland honeyeater defends feeding territories

containing nectar-producing Banksia owers. In general, birds
hold smaller territories in areas with a high density of Banksia
ower clusters than where these are sparsely distributed. The
nectar resource varies over time. When it is scarce, New Holland
honeyeaters exhibit more aggression toward intruders in an
effort to keep them off their patch, but when the sugar-rich liquid
becomes abundant they may relax their hold on territories and
become less aggressive since
there is enough food to go
around. During the breeding
season, pairs of New Holland
honeyeaters also hold breeding
territories. The female is more
involved in nest building and
incubating of eggs and tends
to feed closer to the nest,
while the male defends the
nest but ranges farther aeld
often to the edge of their
territoryto feed.
Phylidonyris novaehollandiae

a 7 in (18 cm)
b Meadows, woodland, and yards

across southern Australia.

c Black-and-white bird with yellow

wing patch and yellow margin on tail.

An intruders red breast
triggers territorial defense
behavior in a resident
male robin. A warning
song can quickly escalate
into a vicious fight in
which one or other of the
opponents might die.

Seasonal variation
European robin


The European robin maintains

territories in both the breeding
and nonbreeding seasons.
Spring and summer breeding
territories average 1.3 acres
(0.55 hectares), while winter territories may
be only half that size. The rst territories to
become colonized in an area are the most
resource-rich, wooded habitats, which are
favored for breeding. During winter,
migrant birds may arrive and colonize
territories in less valuable shrub habitats.
During the breeding season, males sing
and patrol their territories more often;
they are also quicker to attack neighboring
males, which are more likely to trespass.




Male and female robins hold
separate territories in winter.
By the time spring comes around,
some birds have died, permitting
others to increase the size of
their territories. By the following
winter, the territories have been
recolonized by young birds.


Erithacus rubecula

a 41 2 51 2 in (1214 cm)
b Woodland, parks, yards, and farm

hedgerows across Europe.

c Songbird with bright orange-red

breast and face, gray-brown upperparts,
and off-white belly.
d 460

Fixed range
Koalas are mainly solitary animals that occupy distinct home
ranges. In wetter forests in the south of their distribution, koalas
require only small home ranges of 1.27.4 acres (0.53 hectares)
to meet their needs, while in drier, less productive areas in the
west their home ranges can be as large as 250 acres
(100 hectares). A dominant males range
may overlap with those of up to nine females.
Adult males travel widely at night during
the breeding season (October to February),
ghting with rival males and mating with
receptive females. They also call to
advertise their presence and scent-mark
tree trunks using a gland on their sternum.
Phascolarctos cinereus

a Up to 31 in (78 cm)
b Native to eucalyptus forests of eastern Australia.

Introduced to western Australia and nearby islands.

c Woolly marsupial with gray-brown fur on back,

white chin and chest, and long tufts of white hair on
ears. Males are generally larger than females.
d 191
Koalas feed on the leaves of around 30 species of
eucalyptus. Their opposable digits and curved
claws help them grip branches.


A study in New South Wales has
shown that carefully managed
logging can minimize the effect
on koalas home ranges. About
a quarter of the white cypress
pines, a tree that is used as a
daytime shelter by the koala, were
removed from a forest during the
study, but the koalas main food
trees, three species of eucalyptus,
were deliberately left untouched.
The koalas were unaffected by
the loss of the trees.


Controlled supply
New Holland honeyeater


Tigers are generally solitary, so when a stranger is
encountered on anothers territory a fight usually
ensues. They are powerful animals with an
impressive weaponry of sharp teeth and claws.

Both male and female tigers maintain territories. These need to

have dense vegetation for cover, a source of water, and plenty of
prey. Also, a territory usually contains several dens where the tiger
can rest and where females give birth to and rear their cubs.
Territories of male tigers are up to three times larger than females
territories, and male and female ranges overlap. A male tends to
retain exclusive breeding rights to the females in his territory as
long as he can defend them from invaders. To minimize the risk
of conict, tigers leave signs that an area is occupied by scentmarking their territories with urine and feces or leaving scratch
marks on trees. They are also quick to colonize ranges that have
been vacated by another tiger dying. If a male takes over anothers
territory, he may kill any offspring he nds before mating with the
resident females. Humans threaten tiger populations by poaching,
fragmenting their habitats, and reducing the density of their prey.
Tigers hunt by stealth,
approaching potential
prey (which includes
deer, buffaloes, and
wild pigs) silently in
a crouching position,
before launching an
ambush. They kill
by breaking the animals
necks or by suffocation.


Tigers spray urine mixed with a musky
scent from an anal gland onto trees and
rocks to mark their territories. Females
increase their rate of spraying just
before they are ready to mate.

Panthera tigris

a 61 2 1214 ft (23.7 m)
b Forest in northeast China, Korea,

Russia, northern India, and Nepal.

c Orange-red coat with black stripes

and paler underside. Strong shoulders
and limbs with broad paws and long,
retractable claws.

d 375


The estimated
number of tigers
remaining in the wild in India.












In areas of low prey density, such as eastern

Russia, tigers have large home ranges. By contrast,
where prey density is high, as in Chitwan National
Park, Nepal, home ranges are small.


900 1,000


Cat fight


Treetop territory
Brown-throated three-toed sloth

Making their mark

Garnetts galago

Brown-throated three-toed sloths spend most of their time high

in the canopy of rain forest trees, feeding on leaves, sleeping, and
resting. They typically live 6698 ft (2030 m) off the ground and
transfer from one tree to another by using overlapping branches
and climbing plants such as lianas. Sloths move slowly, hanging
upside down from their long limbs, which are tipped with three
hooklike claws. They also metabolize their food slowly and have
relatively small muscle mass, so they are not able to generate
much of their own body heat. Consequently, they seek or stay
out of the sun during the day to adjust their body temperature as
required. Brown-throated three-toed sloths are usually solitary and
occupy a range of less than 5 acres (2 hectares). When ready to
mate, females attract males into their territories by a screaming
call. They have one baby a year, which they carry around on their
chests until their offspring
is able to be independent.

Garnetts galago, also known as Garnetts

greater bushbaby, is a nocturnal primate
that eats fruit, gum, and insects. It is
generally solitary, although it may
encounter other animals when feeding and
females may sleep with their offspring.
Like many other bushbabies, Garnetts
galago has a habit of washing its hands
and feet with urine. It is not clear whether
this behavior is concerned with scentmarking territory, or if it improves the
animals grip when leaping from tree to
tree. Dominant males mark trees more
often than females and subordinate males,
by rubbing branches with a gland on their
chest or with their urogenital region.
Garnetts galagos also communicate by
sound, by vocalizing and by rubbing their
feet against a surface to make a noise.

Bradypus variegatus

a 161 2 31 in (4280 cm)

b Lowland rain forest in Central and
South America.

c Shaggy, coarse, brown-gray coat,

often tinged green with algae. Brown

throat, dark brown forehead, and stripes
on either side of the eyes.

Otolemur garnettii

a 101 2 in (27 cm)

b Coastal, riverine, and highland forests of East Africa,

from Somalia to Tanzania (including offshore islands).

c Coat color varies according to subspecies, from

reddish to gray-brown, sometimes with greenish tones.

d 311
The brown-throated three-toed sloth
moves to the forest floor about once
a week to defecate in a small hole
that it digs with its tail. Sloths move
awkwardly on the ground, but, if
necessary, they can swim across
rivers or areas of flooded forest.


Each day between 40 and 100 forest elephants
visit the Dzanga Bai in the Central African
Republic. Forest elephant herds gather at bais
(swampy forest clearings) to dig for and eat
mineral salts contained in the soil and to drink
the mineral-rich water.

Galagos have huge, lightsensitive eyes that enable
them to see in the dark.
Their large ears can move
independently of one another,
to help them detect noises
made by insect prey.

The home ranges
of dominant male
galagos overlap with
those of several
dominant females
and their young.
Interactions in these
areas are generally
friendly. Transient
males may not have a
specific home range.

Breeding males
Breeding females
large, rounded ears

spherical eyes

The red fox is a very adaptable carnivore that lives in many

different habitats across its extensive range. In recent decades,
it has become an increasingly common xture in urban areas,
where it exists in close proximity to humans. Foxes form small
social groups consisting of a male (known as a dog fox), several
females (vixens), and their dependent young (cubs). Each groups
territory contains a den, or earth, where the females give birth to
and rear their cubs. Well-trodden pathways connect the earth to
other dens, hunting grounds, and food caches. Fox home ranges
vary in size from 25 to 12,350 acres (105,000 hectares)
depending on habitat quality. Territories are smaller and less
stable in urban than in rural areas, probably because the urban
environment changes more rapidly than the countryside.


Town and country

Red fox


Foxes mark their territories with urine
and scent. When a male ignores these
warnings and enters a rivals area,
a fight may ensue, during which the
foxes rear up and attempt to push
one another over.

Vulpes vulpes

a 1835 in (4690 cm)

b Wide variety of habitats in the Northern Hemisphere,
including forest, tundra, desert, farmland, and towns.

c Three color forms exist: one has a bright orange-red

coat with black legs and a whitish underside; the second

varies from silver to black; the third is a mixture.

Foxes frequently scavenge from trash cans in urban
areas. They may also be fed by humans putting
food out in their yards. In some areas, 50 percent
of a foxs diet may be from human sources, and
these foxes will have smaller territories.

Trunk roads
Forest elephant
African forest elephants live in small family groups of between
ve and eight individuals deep in the forests of the Congo Basin.
The dense vegetation makes these animals difcult to study, but
satellite radio tracking indicates their home ranges could cover
approximately 1,243 square miles (2,000 square km). An elephant
may travel 2 3 914 miles (115 km) a day, feeding on grasses and
leaves in the dry season and fruit in the wet season. Forest
elephants visit watering holes daily, not only for water but also
for soil containing minerals, such as calcium, potassium, and
magnesium, which they need to stay healthy. In 2001, genetic
analysis of the DNA of poached elephant ivory revealed that the
African forest elephant might actually be a species distinct from
the African savanna elephant, rather than a subspecies of it.

Generations of forest
elephants have created a
network of trails throughout
the forest, linking fruit trees.
Loxodonta cyclotis

a 814 ft (2.5 m) (male); 7 ft (2.1 m) (female)

b Equatorial forests of Central and West
Africa, in particular the Congo Basin.

c Smaller than African savanna elephant

with rounded ears, straight tusks with pink

tinge, dark skin, and long, narrow jaw.


Forest elephants move along three main
types of highway: boulevards, which
allow them to travel rapidly between
distant, favored areas such as forest
clearings; foraging paths through
medium-density forests with plenty of
food; and alleys around the clearings.


Migration is the periodic movement of animals to and from different areas, usually along
well-dened routes. In long-lived species, individuals make repeated annual migrations
throughout their lives once they are sexually mature. In short-lived species, such as the
monarch buttery, which produce several generations within a season, the migration is
undertaken by a succession of different individuals as the butteries breed along the route.

Why migrate?


Many animals migrate to nd the best location in which

to lay eggs or rear young. This is often related to the
need to avoid the lack of seasonal of food in one area
and exploit an abundance of food elsewhere. Such
movements are usually in response to predictable
changes in the environment, for example, the different
seasons in temperate latitudes, where weather
conditions vary from harsh to favorable, and the
wet and dry seasons in the tropics.





Longest round trip

Sooty shearwater

40,400 miles (65,000 km)

Longest nonstop flight

Bar-tailed godwit

7,200 miles (11,570 km)

Highest journey

Bar-headed goose

33,380 ft (10,175 m)

Longest aquatic journey

Gray whale

12,400 miles (20,000 km)

Largest land migration

Blue wildebeest

1.3 million

Largest air migration

Desert locust

69 billion


Blue wildebeest (inset)
Arctic tern
Atlantic salmon
Gray whale
European eel
Monarch buttery

Barn swallow









Migrations occur all around the world:
by air, over land, and in the oceans.
Many movements are from north to south
and back again with the changing seasons.
Locust migrations are occasional and
cover broad areas in search of food.


Partial and full migrants

Gulf of


500 m




Fox sparrows migrate
along the western
seaboard of North
America. Populations
that breed in Alaska and
Canada migrate south
in fall to overwinter in
the US, leapfrogging
populations with shorter
migrations and those
that do not migrate at all.

In some species all individuals migrate (full migrants),

while in others some remain resident in parts of the
range (partial migrants). Whether or not an animal
migrates can depend on local climate. For example, in
Finland most European robins migrate south to escape
the harsh winter, whereas in the British Isles, where
winters are milder, most robins remain all year round.
Migration can also depend on the stage an animal
has reached in its life cycle. An immature American or
European eel, for example, has no need to migrate to
breeding grounds; it will make the journey from inland
waterways to the Sargasso Sea when it is ready to
reproduce (see p.152). Migration may also depend on
an animals circumstances. Older, more experienced
Eurasian blackbirds in possession of a territory remain
resident while younger, less experienced birds migrate.

Population A
Population B
Population C
Population D

Migrations are often long and hazardous journeys.

Some birds choose to travel over land even though
it is a longer journey, while others go over the sea by
the shortest route, where there are fewer predators
and they can often benet from tailwinds. But birds
migrating over open sea risk winds blowing them off
course (resulting in birds stopping in places they are not
normally seen) or storms forcing them to land on water.
There may be increased risk of predation at the end of
the trip: in fall, the Eleonoras falcon preys on exhausted
songbirds that have own over the Mediterranean to
North Africa. Migrating over land can have its hazards,
too, such as crossing inhospitable places like deserts.
During their annual
migration, Burchells
zebras must risk
crossing the crocodileinfested waters of
the Mara River in
Kenyas Masai Mara
game reserve.

Triggers to migrate include lengthening days as spring approaches,

increased reproductive hormones and fat deposits, and growing
restlessness. Many animals have an innate, near-annual rhythm
that tells them when to migrate. Once in good condition,
favorable weather may be the nal trigger to move. Physiological
changes are often necessary before migration. Birds heart, ight,
and skeletal muscles enlarge, while other organs may diminish in size.
Fishes moving between fresh and saltwater display
changes in their levels of salt tolerance, while
amphibians that move between terrestrial and
aquatic habitats alter the permeability of their skin.




Birds that cross deserts or open oceans cannot
guarantee regular food during migration. They
deposit fat in their bodies as fuel for their flight.
Small birds can double their weight before leaving.

Animals that migrate use various cues to help them
navigate. Compass cues indicate direction and might
include the sun, stars, or the Earths magnetic eld
(see p.131). Visual cues or landmarks, such as
coastlines and mountain ranges, are used
by animals to pilot their way toward
their goal. Distinctive odors are
also used to home in on
breeding or roosting sites.
True navigation relies on a
mental map to determine
position relative to a destination. In some
species, young birds migrating for the rst
time travel with adults in ocks, but in many
the adults leave rst and the young follow
later. They are born with the information for
the distance and direction of their journey.
magnetite around a birds
upper mandible may enable it to
sense the direction of magnetic north

Flying in a V-shaped formation allows migrating
geese to save energy and communicate about
orientation cues. Barnacle geese migrate
seasonally between the high Arctic and more
southerly latitudes.


Birds use a combination of
cues during migration and may
switch from one to another depending
on conditions. If the sky is cloudy, they
may rely more heavily on the magnetic field.

Human-assisted movement
Throughout history, humans have moved animals
around the world for various purposes, sometimes
with disastrous consequences. Goats were
introduced to many oceanic islands to provide food
for passing ships. In many cases, they have since
caused signicant environmental damage by
stripping the land bare of vegetation. People have
also taken animals, such as cats, to new places as
pets, where they then kill the local fauna. And many
species have been introduced in an effort to control
others. For example, cane toads were imported to
Australia to eat pests of sugar cane, but instead
they eat almost anything else (see p.284).



The black rat has colonized
much of the world by stowing
away onboard ships. Other
invasive species have been
transported in the water in
ships ballast tanks or
attached to the hulls.

The opening of the Suez Canal in 1869

connected the northern end of the Red Sea
to the Mediterranean Sea. For a long time,
part of the canal, the Bitter Lakes, was so
salty that few animals could survive in it.
However, over time the salinity has gradually
decreased and Red Sea species have been
able to travel northward into the eastern
Mediterranean. This movement is known as
Lessepsian migration after the engineer of
the Suez Canal, Ferdinand de Lesseps. At
least 300 species are known to have made
this human-assisted migration.

eyesight provides
visual clues to


Triggers and preparation


Ocean bound
Red crab

Once they reach the beach, the crabs head
straight into the ocean to replenish the water
and salts lost during their journey.

For most of the year, the red crabs on

Christmas Island reside in burrows in the
rain forest oor, feeding on leaf litter, but as
the monsoon rains start in November, tens
of millions of these vibrant crustaceans start
marching toward the coast. Their trip
takes about a week; on arrival, males dig
burrows in the beach terraces where they
mate with females (the burrows protect
the crabs and their eggs from heat,
dehydration, and predators). The male
crabs leave soon after to trek back inland,
but the females stay at the coast for two
weeks while their eggs develop. Then, at
the turn of the high tide in the last quarter
of the moon, the females release their
eggs, up to 100,000 each, into the sea.
Gecarcoidea natalis

a 41 2 in (12 cm)
b Forests of Christmas Island and the Cocos Islands,

Indian Ocean.

c Bright red crab with broad, rounded carapace. Males

are usually larger than females.

Across the sea floor

Spiny lobster
In late October and early November,
regimented lines of spiny lobsters make
their way across the sandy bottom of the
Caribbean Sea. The migrations are thought
to be triggered by fall storms that make the
shallow waters cooler and more turbulent.
The lobsters walk mainly at night and take
shelter in crevices or in stationary groups


Each year, thousands of red
crabs are killed while crossing
streets to reach the sea.
The people of Christmas
Island try to reduce
this number by
closing streets at
peak times on the
main migration
paths. Other streets
have safe crossing points
beneath them. Crabs are also
threatened by the clearing of
their forest habitat for phosphatemining and human settlement,
which forces them to travel longer
distances in the open. Fortunately,
63 percent of the island is now
protected by a national park.

47 million

The number of red

crabs that migrate from the forests
of Christmas Island down to the sea
to spawn each year.

in the open during the day. They take

several days to travel 1831 miles
(3050 km) to reach the edge of the deep
ocean channels, where they spread out
along the less disturbed reefs on the
ocean fringe. Females spawn in this
deeper water in spring and early summer,
before returning to the shallows. Once the
eggs have hatched, the larval lobsters drift
on the ocean currents, eventually being
brought back to the Caribbean Sea,
where they mature.

Panulirus argus

a 231 2 in (60 cm)

b Coral reefs and sea-grass beds in the

Atlantic Ocean, Caribbean Sea, and Gulf

of Mexico.

sharp spine
tail tucked
under body


c Brown-gray lobster with yellow spots

on the segmented abdomen.
The spiny lobster is protected from most
predators by the sharp spines on its
exoskeleton, but if attacked it tucks its
tail under its body.

small claw

Each lobster uses its antennae and front legs to
keep in touch with the abdomen and tail fan of the
lobster in front. This in line behavior may be a
defense against predators and also reduces drag.

Anax junius

a 234 314 in (78 cm)

b Still and slow-moving freshwater in North and South

America, the Caribbean Islands, and Asia.

c Dragonfly with distinctive green thorax and two pairs

of large, transparent wings.

North and south

Green darner

Searching for food

Peachpotato aphid

Each spring, a new generation of green

darner dragonies migrates from the
southern United States to the north and
to Canada, where they spend the summer
breeding and feeding on mosquitoes.
Approaching cold fronts in fall trigger the
next generation of dragonies to return
south, sometimes covering as much as
87 miles (140 km) in a day. On arrival,
the dragonies breed and their offspring
overwinter as nymphs, then metamorphose
into adults in spring to start the cycle
again. In this way, successive generations
of offspring migrate rst north then south,
then north again. The migrating dragonies
are a temporary food source for kestrels.

As its name suggests, the peachpotato

aphid moves between different host plants
as an aid to its survival. In fall, adult aphids
mate and lay eggs on peach and related
trees such as apricot and plum. The eggs
remain dormant until spring, when they
hatch into nymphs that feed on the trees
owers, leaves, and stems. Winged forms
of the aphid leave in search of new food on
a wide variety of summer vegetable crops
including potatoes. The female aphids
breed fast, giving birth to live young by
parthenogenesis (see p.328). In fall, the
aphids return to the peach trees. The
peachpotato aphid is a major pest as
a carrier of plant viruses.

Myzus persicae

a Up to 116 in (2 mm) (wingless form)

b Worldwide, on peach trees during fall, winter, and

spring and various crop plants in summer.

c Winged adults have pale green-yellow abdomens

with black heads and thoraxes. Wingless adults are more
variable in color.

Millions of monarch butterflies undertake
the epic journey south to the highlands of
Mexico every fall. Stored fat fuels their
flight, and they may glide on air currents to
save energy.

Suitable conditions for overwintering monarch
butterflies in Mexico occur only in a relatively small
area in the state of Michoacn. The fir trees are
festooned with the orange and black butterflies
from November to March.

One of the most conspicuous of all insect

migrations is that of monarch butteries
in North America. In fall, they move south
to avoid the cold temperatures of more
northerly parts of the continent. They
overwinter in milder climates in a state of
reproductive diapause, meaning they do
not breed during this time. In spring, these
striking butteries move north in search
of their food plant, milkweed (plants of the
genus Asclepias, which produce milky
sap). As they move, females lay eggs and
die, and the new generations continue the
journey. By the time they have reached
the most northerly parts of their range, the
butteries will be second-, third-, or even
fourth-generation descendants of those
that left the south. Monarch buttery

migration appears to be triggered by

changes in day length and temperature,
and there must be a genetic component
to the migration that allows ight routes
to be inherited by offspring, since no
individual buttery ever makes the journey
twice. The overwintering populations of
monarch butteries in Mexico are threatened
by destruction of their forest refuges for
lumber; intact forest is vital to maintain the
microclimate needed for the butteries
survival. Gaps in the forest cover can leave
the butteries susceptible to cold and rain.
Several sanctuaries have been established
for the protection of monarch butteries.
Danaus plexippus

a Wingspan 31 4 41 2 in (8.512 cm)

b Native to open habitats and forests in North America.

Also found in Australasia and parts of Europe.

c Adults have orange wings with black borders and

veins and some white markings. Caterpillars are striped.

Summer range
Spring range
Winter range
Northerly migration

Monarch butterflies that overwinter
in coastal California spend the
summer inland, to the west of the
Rocky Mountains. Those that winter in
the highlands of Mexico travel via Texas
to northern states east of the Rockies,
although some butterflies have been
tracked flying to states to the west.
The distance between summer and
winter ranges can be as much as
3,000 miles (4,800 km).



e r n li mit of m il
k w e ed

Rocky Moun ains

Mass migration
Monarch buttery





500 miles

Like the European sea sturgeon, salmon

are anadromous, meaning that they live
mostly in the sea but return to freshwater
to breed. Chum are one of five species
of salmon that frequent the North Pacific
Ocean and the rivers that border it in
countries including Canada, Japan, Korea,
and the United States. After spending
between one and three years at sea, chum
salmon travel inland as far as 2,000 miles
(3,200 km) up the Yukon River through
Alaska into Canada. The annual salmon
run occurs during fall, with spawning
taking place between November and
January. Around two weeks later, the
adults die, contributing valuable nutrients
to the ecosystem. Their eggs remain
protected in the gravel riverbed over
winter and hatch in early spring. The fry
stay in the river for a year or more before
traveling downstream to the sea between
March and July.
Oncorhynchus keta

a Up to 3 ft 3 in (1 m)
b Temperate and cold waters of North Pacific and

adjacent rivers.

c Silver bluish green salmon. Spawning males develop

darker markings with vertical bars of olive and grayish
red on their sides.

migration hazard
Along their way upstream, migrating salmon must
contend with a variety of dangers, not least hungry
grizzly bears that gather at waterfalls and may wait
for hours to catch a flying meal.


parasitic lice
Salmon is a popular food fish, which has
prompted a boom in commercial salmon
farming. However, the industry is threatening
wild salmon populations by exposing them
to parasites. Sea lice are crustaceans that
occur naturally on the salmons skin, but the
raising of thousands of adult fish in close
proximity in pens causes parasite numbers
to rise far beyond normal levels found in
the wild, lowering the health of the fish.
Young wild salmon migrating downstream
from their spawning grounds pass the salmon
farms on their way to the ocean and become
infested with parasites as they go.

leaping falls
It is hard work returning to the spawning grounds.
Salmon must jump up waterfalls and rapids,
propelling themselves forward against the flow of
water with their strong muscles and flicking tails.

spawning grounds
By the time they reach the spawning grounds, these
sockeye salmon (Oncorhynchus nerka) have
developed distinctive coloration, with a red upper
body and green head. The males have hooked snouts.

fertilizing eggs
Female salmon lay between 2,000 and 6,000 eggs in
shallow depressions called rudds. Males then release
sperm over the eggs before the females cover them
with gravel for protection.

next generation
Recently hatched salmon are known as alevins.
They remain in the rudd for around 12 weeks, until
the yolk sac attached to their belly is used up and
they have to emerge to feed themselves.

153 migration

heading upstream
Chum salmon

Living space 152

occasional migration
Painted lady

Riding currents
Blue shark

The painted lady is a sporadic migrant: in

some years, streams of butterflies head
north from their winter ranges in the
deserts of North Africa, southwest US,
and Mexico; in other years, very few
butterflies are seen. The mass migrations
are precipitated by rains in arid areas that
cause an abundance of food plants for the
butterflies and a consequential boom in
their population. Particularly spectacular
migrations tend to occur in El Nio years.
When their food supply is exhausted, the
butterflies travel north in search of further
food, across the sea if necessary, reaching
northern Europe and the northern US by
midsummer. The butterflies are aided on
their travels by southerly winds.

During the course of a year, blue sharks

in the North Atlantic Ocean undertake
a clockwise migration, riding a series of
ocean currents called the North Atlantic
Gyre. The Gulf Stream takes the animals
up the coast of North America, then they
follow the North Atlantic Current across
to European shores. The Canary Current
carries them to North African waters,
from where the North Equatorial Current
delivers them back to the north coast of
South America and the Caribbean. The
longest distance so far recorded from
a recaptured tagged shark is 4,250 miles
(6,840 km) from Ireland to Venezuela, but
journeys of around 1,900 miles (3,000 km)
are common. The majority of
transatlantic sharks are female. They
mate in the western North Atlantic
and give birth to their pups in the
waters off Spain and Portugal
and in the Mediterranean Sea.

Vanessa cardui

a Wingspan 23 in (57.5 cm)

b Meadows, hillsides, and marsh habitats worldwide,

except for Antarctica.

c Butterfly with orange and black upper wings with

white spots on pointed black tips of the forewings.

Moving in convoy
Boree moth
Caterpillars of the boree moth have
voracious appetites, which compel them
to keep moving. Having stripped bare
the tree they were living on, they have
no choice but to venture forth in search
of a new food source. As each caterpillar
walks, it extrudes a trail of silk, which is
faithfully followed by its siblings, creating
the illusion of one incredibly long, hairy
caterpillar snaking across the ground.
The caterpillars nest communally in their
food tree in a ball, surrounded by silk and
shed hairs, which has given rise to their
alternative name of the bag-shelter moth.

Prionace glauca

a Up to 13 ft (4 m)
b Tropical, subtropical, and temperate waters


c Long-snouted fish with long pectoral fins. Dark

indigo-blue upper side shades to white underneath.
Weighs up to 440 lb (200 kg).

tracking sharks
Since the early 1960s, blue sharks have
been tagged to provide information on their
movements. Anglers and commercial
fishermen are encouraged to take part in
the program, applying identification tags to
sharks they have caught, either intentionally
or accidentally, before they are released.
The tags are attached to the sharks dorsal
fin or implanted in the muscle of its back. If
the shark is recaptured, scientists can start
to draw a map of its migratory journeys.
More recently, satellite tags have allowed
the sharks travels to be mapped without
the need for their recapture.
on the Move
The migrations of the Atlantic population of blue
sharks have been the most extensively studied,
but the Pacific population has been found to
migrate long distances too. There is also
a population in the Indian Ocean.

Ochrogaster lunifer

a 11 2 in (4 cm) (mature caterpillar); 11 2 in (4 cm)

(moth wingspan)
b Acacia and eucalyptus trees in Australia.
c Caterpillar is gray and hairy with a brown head. Adult

moth is dark gray or brown with a pale dot in the center

of the forewing.

Return journey
European eel

Seeking freshwater
European sea sturgeon
The European sea sturgeon is anadromous, spending much of its
early life at sea and returning to the river of its birth when sexually
mature. Females lay up to 6 million sticky eggs in shallow water,
on gravely riverbeds. After hatching, juveniles make their way
downstream, becoming acclimatized to the increasing salinity
as they near the sea. The European sea sturgeon was previously
abundant around Europe, but it is now one of the most
endangered fish as a result of over-exploitation for its eggs (which
are eaten as the delicacy caviar), the deteriorating quality of its
freshwater spawning grounds,
and obstacles to migration,
Acipenser sturio
such as hydroelectricity
a Up to 161 2 ft (5 m)
projects. This sturgeon is now
b Coastal waters of northeastern
thought to breed in only one
Atlantic, Mediterranean, and Black Sea,
and adjacent rivers.
river basin in France, and a
captive breeding program
c Olive-black upper body with white
underside. Five rows of scutes.
has been established.

Unlike the European sea sturgeon, the

European eel spends most of its life in
freshwater but returns to saltwater to
breed, a behavior known as catadromy.
Having started life in the ocean, young eels
(elvers) arrive at estuaries and travel inland
along rivers. The eels remain upstream for
more than ten years, feeding and maturing.
In July each year, adult eels return to the
sea, sometimes crossing wet grass to
reach a different watercourse. These eels
develop larger eyes to see in the ocean
depths and a silver coloration below that
helps conceal them from predators.

Anguilla anguilla

a Up to 414 ft (1.3 m)
b Sargassum (seaweed) beds of the western Atlantic,

coasts of the northeastern Atlantic, and adjacent rivers.

c Long, narrow-bodied fish with continuous dorsal,

anal, and tail fin. Dark scales on top and yellow below.

Young and old

The European eel
progresses through
various stages. The firststage larva is called a
leptocephalus (left). It
then becomes a
glass eel, an elver,
and a yellow
eel, before
becoming a
fully mature
adult (silver
eel, below).


finding the spawning grounds

From 1904 to 1922, Danish biologist
Johannes Schmidt led expeditions in the
Mediterranean Sea and the North Atlantic to
find the spawning grounds of the European eel.
He recorded the length of larval eels caught in
various locations, eventually catching the
smallest, 3 8 in (1 cm) long, in the Sargasso
Sea. In the western Sargasso, Schmidt also
found the breeding grounds of the American
eel (Anguilla rostrata ), a related species that
matures in the rivers of the eastern United
States. The larvae drift on ocean currents to
reach coastal waters, the American eel larvae
exiting the current before the European eels.

larvae size


13/4 in
1 in


/5 in
/8 in


larvae size


1,000 miles


Dangerous journey
European common toad

Back to the birth pool

Yellow-spotted salamander
Yellow-spotted salamanders hibernate in woodland, beneath leaf
litter or underground; with the rst thaws of snow, they migrate to
their breeding pools. This can be as early as December to February
in southern parts of their range or as late as March to April in the
north. They return to the pool in which they were born, which is
usually less than 650 ft (200 m) from their hibernation areas. After
breeding, the adults leave the eggs
Ambystoma maculatum
to develop into larvae, which may
a 41 2 10 in (1125 cm)
overwinter in the pools in their larval
b Moist mature woodland in
form or metamorphose into adults
and leave the pools that year.
Occasionally, these salamanders
will lay their eggs beneath the ice
of a frozen pool.

Like salamanders, European common toads attempt to migrate

back to the pond in which they were born in order to breed. The
migration begins in fall but is interrupted by a period of hibernation
in midwinter, when the toads hole up underground to wait until the
cold weather has passed. They resume their journey in spring, and
it is then that large numbers of toads are frequently killed while
crossing streets built on their traditional migration routes. Those
toads lucky enough to reach the breeding ponds mate in a frenzy
between March and June, before
returning to their home range, which
may be anything between 180 and
5,250 ft (55 and 1,600 m) away from
the pond.
Bufo bufo
a Up to 7 in (18 cm)
b Terrestrial and wetland habitats throughout

North America.

most of Europe.

c Stocky, dark gray, black, or

brown salamander with two rows of
bright yellow spots along the sides
of its body.

c Broad, squat-bodied toad with warty olive

green-brown skin. Exudes a distasteful secretion
to ward off predators.

Long-distance swim
Leatherback turtle
This giant of the ocean makes lengthy
annual migrations. Using ocean currents
to help it on its way, the leatherback turtle
feeds in the rich, temperate waters of the
northern and southern Atlantic and Pacic
oceans in spring and summer, but it
returns to more tropical climates to nd
beaches on which to lay its eggs. Although
only females venture onto land, males also

d 284, 370

migrate, coming close to shore to mate

with the females. Satellite tagging has
revealed extensive migrations: one recordsetting leatherback was tracked swimming
12,774 miles (20,557 km), from a beach in
Indonesia across the Pacic to the west
coast of the US and part of the way back
again. Leatherbacks are prone to eating
plastic refuse, mistaking it for jellysh, and
can choke to death as a result. They are
also sometimes caught in shing nets.
Dermochelys coriacea
a 514 7 ft (1.62.1 m)
b Temperate, subtropical, and tropical waters


c Largest of the marine turtles, with relatively long

front flippers.

With a span of up to 9 ft (2.7 m),
leatherback turtles have the
longest front flippers relative
to their size of any turtle.
They usually cruise at around
114 miles (2 km) per hour.

Tunnels or underpasses built beneath roads
close to breeding ponds can help reduce
toad mortality from collisions with cars.
Other measures include warning signs and
people carrying the toads across in buckets.


Scientists increasingly use satellite tags
to track the movements of leatherback
turtles in the worlds oceans. The tags are
mounted on harnesses that are designed to
disintegrate gradually and fall off the turtle.
They are usually xed to the females once
they have nished nesting and to males that
are accidentally caught at sea by shermen.
For up to two years, a tag can relay
information about a turtles whereabouts
via satellite to a computer. In addition to
location information, the tag transmits data
such as dive depth and duration, which give
clues about behavior such as foraging.

Female leatherback
turtles lay an average
of 110 eggs per clutch
in a hole in soft sand
that they dig with their
flippers. The young
hatch after 60 to 70
days and rush headlong
to the sea.

The females pull themselves onto gently sloping,
sandy, tropical beaches to lay their eggs above
the high-tide line. Most species of turtle return to
the beach of their birth and the leatherback is no
exception, although they may visit another beach
in the same general vicinity.


0 km
(0 miles)
Plateau of Tibet
Breeding ground

Mt Everest

Bar-headed geese fly over the highest

mountain range in the world when migrating.
The greatest height they have been recorded
flying at is 33,380 ft (10,175 m) above
sea level.




Bar-headed geese reach extreme high

altitudes during their migration over the
peaks of the Himalayas. The birds are
supremely adapted to the demands of their
trip: their wings have a large surface area
relative to the birds weight to provide extra
lift; their blood hemoglobin absorbs

oxygen faster than that of other birds; and

their downy feathers keep them warm by
trapping body heat generated by ying.
Bar-headed geese are capable of traveling
more than 1,000 miles (1,600 km) each day
and can y at speeds of 50 mph (80 kph).
Annual migration allows the geese to avoid
severe winter storms on the high plateaus
of Central Asia, where they breed in the
summer, and summer monsoon rains on
the Indian subcontinent.

200 km
(125 miles)

400 km
(250 miles)

600 km
(375 miles)

Anser indicus
a 2832 in (7082 cm)
b Mountain lakes and wetlands in Central Asia.
c Pale gray-bodied goose with orange legs and bill.

800 km
(500 miles)

Two distinctive black bars on the head give it its name.

Bar-headed geese spend the winter in wetlands in India,
northern Myanmar, and Pakistan, feeding on grasses and
cereal crops such as barley, rice, and wheat.

In summer, bar-headed geese live at high altitudes,
on the Tibetan Plateau in Tibet, Qinghai in China,
and Ladakh in Kashmir. The geese, such as the
ones on the water in this photograph, congregate
at mountain lakes to breed and feed on short grass.

Roaming the ocean

Wandering albatross
This albatross spends most of its life at
sea, soaring above the waves or resting on
the surface. It ventures thousands of miles
across the Southern Hemisphere from its
breeding islands close to the Antarctic
Circle, occasionally crossing the equator
or circumnavigating Antarctica in its search
for food. One bird was tracked traveling a
staggering 3,700 miles (6,000 km) in just
12 days. Wandering albatrosses nest on
a few islands covered in tussock grass
between November and July. Males and
females take turns incubating the egg and
caring for their single chick while the other
feeds at sea.
Diomedea exulans
a 31 2 41 2 ft (1.11.4 m)
b Open ocean and remote oceanic islands of the
Southern Ocean.

c Adults have white bodies, with white and black

wings. The large, pink hooked bill has tubular nostrils.


It is estimated that 100,000 albatrosses
drown each year on longlines baited for
bluen tuna. The lines, which are pulled
behind shing boats, can be up to 80 miles
(130 km) long and carry as many as 10,000
hooks. The birds try to catch the bait while
it is close to the surface, become hooked,
and are dragged underwater as the lines
sink. International conservation efforts
advocate the use of measures by shermen
to help prevent these deaths.

Young albatrosses take
up to ten years to reach
maturity and during
that time may hardly
ever make landfall.
They often accompany
fishing vessels, feeding
on discarded fish.

9.8 ft

The wingspan of the

wandering albatross, which
is greater than that of any
other flying bird.


Peak performance
Bar-headed goose


Bar-tailed godwits (seen here in a mixed flock with
other shorebirds) frequent estuaries in the winter.
They use their long, sensitive bills to probe for
crustaceans, mollusks, and worms in the mud.



and ew 6,340 miles (10,219 km) nonstop
to Yalu Jiang, China, arriving on March 24.
She spent ve weeks there before leaving
on May 1 and traveling to Alaska, stopping
occasionally en route. E7 arrived at her
breeding grounds at Manokinak on the
YukonKuskokwim Delta on May 15, where
she stayed until July 17, before moving to
another site in the delta. On August 29, she
departed and smashed her own record by
ying nonstop to New Zealand, a distance of
7,200 miles (11,570 km), in just over 8 days.

In 2007, biologists fastened satellite

transmitters to 13 bar-tailed godwits on their
feeding grounds in New Zealand. They were
about to witness the longest-ever nonstop
ights made by a bird. A female, nicknamed E7,
left the mouth of the Piako River on March 17

identification tag


This large wading bird is a long-distance

globe-trotter, capable of ying nonstop
from breeding ranges in the Arctic regions
of Europe, Asia, and western Alaska
across the equator to feeding grounds in
South Africa, Australia, and New Zealand.
During the Northern Hemisphere summer,
bar-tailed godwits are resident on the
coastal tundra of the far north, where they
lay between two and four eggs in cupshaped nests in the grass. After raising
their young, the birds y to refueling
stations, such as estuaries, to feed and
bulk up their bodies before ying south to

the Southern Hemisphere. As the Southern

Hemisphere winter approaches, the birds
molt into their russet-colored breeding
plumage and put on weight in advance of
their northward migration. Some bar-tailed
godwits overwinter in Europe, where they
are closer to their breeding grounds and
require less energy to get there. Although
their feeding rate is the same as birds that
migrate from Africa, they spend less time
feeding before they begin their journey.
Limosa lapponica

a 141 2 16 in (3741 cm)

b Arctic coastal tundra and sandy intertidal areas

in South Africa, Australia, and New Zealand.

c Neck, breast, and belly are brick-red in summer,

changing to off-white in winter.

115 May
4,500 miles


1724 March
6,340 miles
(10,219 km)

Longest nonstop flight

Bar-tailed godwit




29 August7 September
7,200 miles
(11,570 km)


The bar-tailed godwit
lays down huge fat
reserves (up to 55
percent of its body mass)
before embarking on its
epic nonstop journeys.
Its digestive tract shrinks
in size to make room for
the extra fuel.

The young of many water birds follow their

parents on their rst migration in order
to learn the route. Naturally occurring
whooping cranes migrate 2,500 miles
(4,000 km) between a single summer
breeding area in Wood Buffalo National
Park, Canada, and a single wintering site
in Aransas National Wildlife Refuge,
Texas. The birds reliance on just two sites
prompted conservationists to establish
other populations in Florida and Wisconsin,
and to assist birds in their migration (see
panel, right).


In 2001, conservationists hatched a plan to reintroduce whooping
cranes to Wisconsin. To teach the hand-raised birds the migration
route from Wisconsin to Florida the cranes were imprinted on an
ultralight aircraft. The birds were played the noise of the aircrafts
engine while still inside the egg and soon after hatching were
introduced to the plane. Their rst ights were taken following the
aircraft down the runway and they eventually followed it 1,250 miles
(2,010 km) across seven states to Florida.

Grus americana
In 1941, the migratory flock of whooping cranes numbered just 15
birds. In 2007, there were 73 mating pairs in Wood Buffalo National
Park, 53 resident birds in Florida, and 52 birds in Wisconsin.

Following the sun

Barn swallow
Barn swallows breed throughout the Northern Hemisphere in
summer but travel south to sunnier climes in South America,
Africa, southern Asia, and northern Australia for the northern
winter. They migrate during the day, feeding on insects caught on
the wing. In contrast to some migratory birds, they do not put on
much weight before the journey. This makes them vulnerable
to starvation when crossing large areas with little
food such as the Sahara Desert. They
are also at risk from storms and
exhaustion. Barn swallows can
cover up to 200 miles (320 km)
per day when migrating.

a 414 ft (1.3 m)
b Open, grassy plains and marshland in northern

Canada and Texas, Florida, and Wisconsin.

c Mainly white with black primary feathers and red and

black markings on the head. Long neck and legs.

Barn swallows migrate in large flocks and
frequently stop to rest on structures such as
telegraph wires. Flocks gather in premigratory
roosts in late summer and early fall before
embarking on their southerly journey.

With its long, pointed wings and
elongated tail streamers, the barn swallow
is well-designed for long-distance flying.
Hirundo rustica

a 7 in (18 cm)
b Open grassland, meadows, and farmland habitats

worldwide except Australia and Antarctica.

c Off-white underparts, metallic blue above with

a red-brown throat and forehead.

Once they have found
a good nectar source,
hummingbirds (especially
males) may defend it
from intruders. They
keep a close watch from
a nearby perch on an
exposed branch.

Vertical migration
Long-tailed sylph

Population pressure
Norway lemming

Several species of hummingbird show

altitudinal migration, moving up and
down mountainsides in search of nectarproducing owers. The long-tailed sylph
frequents forests in the Andes at altitudes
of between 3,300 to 9,800 ft (1,000 to
3,000 m). Hummingbirds require an
abundance of energy-rich nectar. They
may move to higher altitudes for a period
of weeks or months when the owers
come into bloom; alternatively, they may
travel each day from lower elevations.
Hummingbirds play an important role
as pollinators in the ecosystem.

Norway lemmings migrate sporadically

every few years when their populations
explode. In a typical year, the rodents live
in tunnels beneath the snow in winter,
moving to higher or lower ground in spring.
There they live on mountain meadows or
in forests, continuing to breed before
returning in fall to the alpine zone. In years
with a mild winter, an early spring, and a
late fall, food is abundant, reproduction is
fast, and survival of litters is high, resulting
in a summer soaring of lemming numbers.
Lemmings are often wrongly said to
commit mass suicide by jumping from
cliffs. Population pressure can trigger
mass migration, usually away from the
meadows toward the forests. Obstacles in
their path, such as boulders, rivers, cliffs,
or ravines, may force the lemmings into a
bottleneck, causing them to panic and
take reckless ight.

Aglaiocercus kingi

a 471 2 in (1019 cm)

b Subtropical and tropical moist montane forest in

Bolivia, Colombia, Ecuador, Peru, and Venezuela.

c Bright iridescent blue and green feathers. Tail is

longer than body.

Lemmus lemmus

a 45 in (1013 cm)
b Tundra, mountain meadows, and alpine and boreal

forests of Scandinavia and Russia.

c Rounded, brown and black rodent with a short tail.

Flattened claw on the first digit of the front feet aids
digging in snow.


Learning the route

Whooping crane

Caribou frequently cross lakes and rivers while
migrating. They are strong swimmers, and their
thick, air-filled coats help them stay buoyant and
warm in the icy water. Both sexes have antlers, but
they are larger and more branched in males. These
caribou are crossing the Kobuk River in Alaska.

Caribou undertake one of the most arduous annual migrations of

any terrestrial mammal. Herds may number thousands of animals
and complete a 3,100-mile (5,000-km) round trip, visiting spring
calving areas and summer and winter feeding grounds. They are
forced to move on by the seasonal availability of the tundra plants
on which they feed. In summer, the herds take refuge from ies
and mosquitoes in windy coastal areas; in winter, they move into
subarctic boreal forests, where snow cover is less than on the
open tundra. Caribou herds have been recorded running as fast
as 50 mph (80 kph) while migrating. Groups are largest during the
spring migration and smaller during fall, when mating occurs.
Caribou are also called reindeer, particularly in Europe and Asia,
where many are partially
domesticated and are tamer
Rangifer tarandus
than wild animals. Reindeer
a 521 2 ft (1.52.3 m)
from herds in Eurasia have
f Arctic tundra and boreal forests of the
US, Canada, Asia, and northern Europe.
been introduced to North
America, where they may
c Coat is brown in summer and gray in
winter, and rump is white.
mix with the native caribou.
Norwegian reindeer
grazing on lichens and
other plants beneath the
snow. The reindeers
broad hooves act like
snowshoes to help them
walk on the snow and to
dig beneath it.

A female caribou tends to
her minutes-old twin calves
in Canadas Northwest
Territories. Twins are rare;
most females have only one
calf each year. Births take
place in May or June on
inland calving grounds.

Although young caribou
can run soon after birth,
large numbers of calves
succumb to predators
such as gray wolves,
which track the migrating
herds and stalk the
birthing grounds looking
for easy prey.



Native peoples of the Arctic and subarctic have a close association
with reindeer, relying on them for food, skins, and transportation.
These people live a nomadic life, moving with the semidomesticated
herds that are making their way between the coast and inland areas.
Reindeer are rarely bred in captivity, but they have been tamed for
milk production and to pull sleighs. Here, a Nenet herder leads her
reindeer across snow-covered pastures in Siberia. Reindeer herding
is also an important part of Sami culture in northern Scandinavia
(Lapland). Native peoples in North America and Greenland have a
long history of hunting wild caribou for their meat and hides.


Endurance test

Like many other bats living in temperate

zones, little brown bats migrate between
several different roosts throughout the
year. On emerging from hibernation in
spring, they travel to summer roost sites,
often in buildings, where the females give
birth to a single pup between May and July.
In fall, bats transfer to hibernation sites,
where they spend winter in a state of torpor.
This behavior helps them survive a time of
year when few insects are available to eat.
Myotis lucifugus

a 214 4 in (610 cm)

b Forested areas of North America, from southern

Alaska and Canada to the highlands of Mexico.

c Small, light to dark brown-colored bat with a plain

nose and relatively short ears.
Migrations to winter sites need not
always be toward warm areas.
Little brown bats may fly north,
sometimes up to 310 miles
(500 km), to spend winter in
a cave or disused mine.

African odyssey
Straw-colored fruit bat
As their name suggests, these African
bats feed on fruitand they undertake
quite a journey to nd enough of it to fulll
their requirements. The bats follow the
annual rains north through sub-Saharan
Africa before returning south at the end
of the rainy season. Their extensive
migration takes them hundreds of miles.
Along the way, they spend the day in large,
noisy colonies in tree roosts. At night, they
leave their roosts and venture forth to nd

dark brown to
black wings

plain nose

Ocean voyagers
Humpback whale

When spy-hopping,
a whale lifts its head
vertically out of the
water while rotating to
look around. It may be
checking for landmarks
while migrating.

Many of the great whales make

extraordinary journeys through the worlds
oceans in search of food and safe places
to give birth, and humpback whales are
no exception. The high-latitude oceans
provide rich feeding grounds during each
hemispheres summer, and here the
whales gorge themselves to store up
energy for their long migration and the
months that follow. In the Northern
Hemisphere, humpback whales feed
more often on sh, whereas those in the
Male escorts accompany female whales in the
breeding grounds, warding off competitors with
assertive behavior such as tail-slapping. They
may also sing complex songs to attract females.

fruit, spreading out to feed. The vast

population of straw-colored fruit bats plays
a vital role in pollinating and dispersing the
seeds of many economically important
plants, including lumber trees such as
iroko, a valuable hardwood, and cash
crops such as cashew nuts, mangoes,
and gs.
Eidolon helvum

a 51 2 81 2 in (1422 cm)
b Forest and savanna in sub-Saharan Africa, the

southern Arabian Peninsula, and Madagascar.

c Dull brown or gray bat with straw-colored fur on its

neck and back. Large eyes and long, narrow wings.

Every November, between 5 and 10 million fruit
bats descend on Kansaka National Park in Zambia.
Their arrival coincides with a boom of fruit in the
park. It is estimated that the bats eat 5,000 tons of
fruit each night.

Southern Hemisphere eat mainly krill. As

winter approaches, humpback whales
move toward the Equator on a journey of
3,100 miles (5,000 km) across the open
ocean in search of warm, sheltered waters,
where the females give birth and mating
takes place. During this time, adults
survive by metabolizing layers of fatty
blubber beneath the skin, and the calves
drink their mothers rich milk. Eventually,
they must return to polar waters to feed.
The population of humpback whales in the
northern Indian Ocean may be resident all
year (see panel, below).
Megaptera novaeangliae

a 4659 ft (1418 m)
b Open oceans worldwide, except extreme north

and south.

c Predominantly black whale with white undersides to

the long narrow flippers. Scalloped edge to fluke.

d 246











Northern Hemisphere humpbacks spend

the summer in feeding grounds in the
northern Atlantic and Pacic oceans. In
winter, they migrate south to breed in
warmer waters around the Caribbean,
West Africa, Japan, Hawaii, and Mexico.
Southern Hemisphere humpbacks follow
the same pattern, spending the summer
feeding in the rich, cold waters of the
Southern Ocean off Antarctica and
migrating north to overwinter in the warm
waters of Australia, the Pacic islands,
southern Africa, or South America.



Seasonal homes
Little brown bat

Major feeding
areas (summer)
Major breeding
areas (winter)
Main migration


Wildebeest calves are born in the middle of the
herd and within days can keep up with the adults
when running. This helps them avoid becoming
prey during the migration.

Mass movement
Blue wildebeest
Each year, around 1.3 million white-bearded
wildebeest (one of ve subspecies of the blue
wildebeest) undertake a spectacular migration
on the plains and savanna grasslands of East
Africa (see map, right). Their herds are further
swelled by hundreds of thousands of other
ungulate species, including Thomsons gazelles,
zebras, and elands. The animals are compelled
to move on in search of fresh food, water, and
minerals, such as phosphorus, that are
important to their health. It is deciency in

this mineral that is thought to spur the herds

to leave their dry-season ranges just before
the onset of the rainy season and head south
to the short-grass plains. Along the way, the
wildebeest have to contend with predators,
including lions and hyenas on the plains and
crocodiles in the rivers.



Dry Season


Connochaetes taurinus

a 614 7 ft (1.92.1 m)
b Plains and savanna grasslands in East Africa, from
the Equator south to the tip of South Africa.

c Large antelope with slate gray to brown coat and

black face, mane, and tail. Curved horns resemble those

of a cow.

Wet Season


50 miles



White-bearded wildebeest follow
a clockwise migration around the
Serengeti Plain. Soon after calving
in the southeastern plains during
the wet season (January and
February), the herds head
northwest, toward Lake Victoria.
By June, they have reached the
transitional pastures. The herds
then turn north toward the Masai
Mara, where they spend the dry
season (July to October) before
Manyara returning south in November to
complete the circle.


A female humpback whale makes the return
journey to polar waters with her young calf. The
calf is weaned at about 11 months but may stay
with its mother for more than a year.



As they cross the Serengeti Plain in Tanzania,
white-bearded wildebeest throw up clouds of red
dust from their hooves. Not all wildebeest are
migratory; smaller herds may remain resident
throughout the year.

Animals produce some of the most impressive architecture on Earth. Their building
methods vary from simply fetching and dropping materials to more advanced
construction techniques, including interlocking, weaving, or gluing materials together
to ensure they stay in place. Some animals make their nests out of pliable substances
such as mud, while others spin structures from silk, or dig burrows in the ground.
Social animals build large structures relative to their body size by working as a team.

Why build?
Animals build structures for a variety of functions.
Many live in their constructions for only a short time; for
example, caddis y larvae build themselves a protective
case from grains of sand or other materials, and some
frogs excavate shelters to help them stay moist during
drought. Animals often build structures for raising
young, as in the case of birds nests and mammals
burrows. Social insects, such as ants, termites,
bees, and wasps, construct elaborate nests
to house entire colonies. Some animals build
structures to capture prey or store food, while
others communicate using architecturefor
example, bowerbirds build courtship stages.
Female polar bears dig dens under the snow to provide
a safe, warm environment during the winter when they
give birth to their cubs. They suckle them for several
months before emerging into the spring sunshine.

Building behavior
Animals may be genetically programmed to build in a
certain way without an image of the end goal and its
function, or they may display a degree of ingenuity and
exibility. For example, a beaver can adapt the shape of
its dam according to local conditions. Building usually
involves relatively simple, repetitive behaviors, and most
animals select or make standardized materials for their
constructions. For example, caddis y larvae reject
sand grains that are either too small or too large for their
cases, and martins choose just the right consistency of
mud for their nests. Most animals use their feet or
mouthparts to manipulate building materials; their level
of specialization depends on whether they are used for
other functions as well, such as feeding.


To nd out how the ability of adult male village weavers to build nests
was affected by their experience when young, researchers gave some
edgling males fresh green building materials to handle (control
group), but other edglings received none (experimental group). When
a year old, the experimental males were unable to weave a single strip
in the rst week that they were given reed grass. After three weeks
of practice, their success rate at weaving was still only 26 percent,
whereas the experienced control group achieved a rate of 62 percent.



Animal architects










90 100


Both weaver ants (right) and tailorbirds (above)
use silk to stitch leaves together to make their
nests. Weaver ants use silk extruded from their
own larvae, while tailorbirds make use of the silk
from spiders webs. They may also use plant fiber
or even stolen household thread. These similar
sewing behaviors have evolved independently
of one another in unrelated species.

The natural world provides numerous organic and mineral building

materials, such as grasses for weaver nests and trees for beaver
lodges, sand and pebbles for many sh nests, and the earth itself
for burrows. Some materials require little in the way of processing,
but others need to be manipulated or mixed with other substances,
such as saliva or water, before being used for building. Many animals
secrete building materials, such as silk, mucus, or wax, from
glands in their bodies. Some use materials derived from others in
their constructions, for example many birds collect soft feathers or
fur to line their nests, whereas others nd spider silk with which to
build them. Animals can be very choosy about the materials they
use in their architecture, selecting particular colors, for example.


Paper wasps chew fibers from dead wood
to make a paper pulp with which they
construct their intricate chambered nest,
which is resistant to water. Male featherfin
cichlids pile sand into a nest that also
serves as an arena for displaying to
prospective mates. Mute swans collect
reeds and other bankside vegetation
to build their nests.





Silkmoth caterpillars spin a cocoon of
silk around themselves for pupation.
Spiders also use silk, but for building webs.
Termites mix saliva with mud to construct
the walls of their termite mounds. Cave
swiftlets also use spittle to build their
nests. Other animal-derived materials
include beeswax and mucus, which is
used by some frogs for foam nests.




Hermit crabs take over the empty shells
of marine mollusks, including the edible
periwinkle. As the crab grows, it moves
into progressively larger shells. Many
animals live in cavities that were created
by another species; for example, the elf
owl nests in old woodpecker holes in the
stems of cacti. Increasingly, humans
provide homes for wildlife, such as a bird
house for a colony of purple martins.





Building materials
A pair of northern flickers
takes between one and two
weeks to excavate a cavity
in a tree, where they lay
their eggs and rear their
chicks. Abandoned flicker
nests create homes for
other cavity nesters.


Stone walls
Hard corals
The worlds most impressive underwater constructions are built
by colonies of tiny invertebrates. Anthozoan coral polyps, most
measuring only a few millimeters across, secrete skeletons of
aragonitea form of calcium carbonateas they grow. Over time,
the skeletons build up into structures known as coral reefs. Coral
reefs are typically found in shallow, clear water around tropical
Corals take different forms according to how the
polyps grow and secrete their skeleton. Over time,
reefs erode and are compacted to form limestone.

coasts or on top of sea mounts (the summits of underwater

volcanoes). This is because corals require abundant sunlight
for their symbiotic algae, zooxanthellae, to be able to
photosynthesize and provide them with nutrients. Corals grow
asexually by budding or division (see p.328). In addition, corals
may reproduce sexually (see p.333). The resulting larvae
eventually settle on the ocean oor and grow into new polyps.

1,240 miles

The length
of the Great Barrier Reef, the
worlds largest reef system



Each polyp sits in a cuplike exoskeleton of
aragonite. Adjacent polyps are linked to one
another by connective tissue. Tentacles
surround the central mouth, through which
food is taken in and waste is ejected. The
mouth leads to the gut cavity. The tentacles
bear stinging cells called nematocysts,
which are used to paralyze or kill prey.


gut cavity

connecting tissue
between adjacent



The 3,000 or so
individual reefs and 900
islands that compose the
Great Barrier Reef can be
seen by satellite. The reef
is located in the Coral
Sea, off the coast of
Queensland, Australia.

Delicate case
Paper nautilus

Diving bell
Water spider

The female paper nautilus (or knobby

argonaut) secretes a paper-thin calcareous
shell that serves as a shelter for the animal
and a brood chamber for its eggs. The
paper nautilus is actually a type of
octopus, only distantly related to the true
Nautilus species, and its shell lacks the
gas-lled chambers found in Nautilus
shells. The paper nautiluss egg case is
usually 41 2 6 in (1215 cm) long
but may reach more than 10 in
(25 cm). It is secreted by
the web between a
pair of the animals
eight arms and is
enlarged along
the outer edge
as the paper

The water spider is able to spend its whole life beneath the waters
surface, thanks to its own supply of air. It spins a web that is
anchored to underwater vegetation, and then lls it by returning
repeatedly from the surface with air bubbles. This diving bell
serves as a place to breathe, hide from predators, consume prey,
and breed. Male water spiders build their diving bells adjacent to
those of females and break through to the adjacent bell when
ready to mate. The female creates a silken cocoon around 3070
eggs at the top of her diving bell. Water spiders may seal their bell
when hibernating and build a
separate bell elsewhere in
Argyroneta aquatica
which to shed their skin.

Argonauta nodosa

a 412 in/1030 cm (female), 114 11 2 in/34 cm (male)

f Surface waters of Southern Hemisphere oceans, from

the Indo-Pacific to the east coast of South America.

c Octopus with tentacles covered with suckers.

Chromatophores in the skin allow it to change color from
red to silver. Only females possess a shell.

The water spider rarely needs to
replenish the air supply in its diving
bell, because oxygen diffuses in and
carbon dioxide diffuses out.

a 14 1 2 in (0.81.5 cm)
f Ponds, streams, ditches, and shallow

lakes in northern and central Europe and

northern Asia.

c Brown-gray above water; takes on a

silvery sheen underwater because air
bubbles are trapped against the body.

Water spiders wait for
prey, such as insects and
small fish, to get close to
their underwater lair.
They swim out and
subdue their prey with
venomous jaws, before
returning to their diving
bell to eat it.

Complex web
Black and yellow
garden spider
The black and yellow garden spider is one
of several species of orb-web spiders,
which weave relatively large, circular webs
that are suspended vertically to catch prey.
The black and yellow garden spider spins
a web that is up to 2 ft (60 cm) in diameter
and positioned 28 ft (60240 cm) above
the ground. Orb-web spiders have three
claws on each foot, to help them handle
the threads while spinning. Silk is extruded

from several spinnerets on the underside

of the spiders abdomen. The spinnerets
are served by different glands that produce
silk of different types and thicknesses.
For example, sticky thread is used for
prey-catching parts of the web. Orb-web
spiders typically consume much of their
web each evening and rebuild it anew
for the next day.
Argiope aurantia

a 3 8 114 in/928 mm (female); 316 3 8 in/ 59 mm (male)

f Sunny habitats, from southern Canada, through the
US, Mexico, and Central America south to Costa Rica.

c Oval abdomen bears striking yellow markings on

black. Black legs with red or yellow near the body.

The spider sits and waits on its web
for an insect to become trapped.
The central, strengthened part of
the web is called the stabilimentum.


The number of
minutes it takes
most orb-web spiders to
build an entire web.
First, the spider connects two structures with a strand
of silk forming a bridge line or primary thread. Then
it begins the web by constructing scaffolding lines,
which connect the structure to its support. Next,
the spider makes a frame of threads radiating from
a central hub. Then it attaches a temporary spiral to
the radial threads. Finally, the web is completed by
the spider spinning close spirals of sticky thread.


spiral of
sticky thread

free zone






turning point



Multipurpose tower block

African mound-building termites
For such small insects, termites construct incredibly
large and elaborate structures in which to live. A
mature termite mound may be home to 35 million
individual termites. Inside, there is a complex
network of chambers, passageways, and
ventilation channels that extend above and
below ground level. The mound has specic
areas designated for waste, for storing and
growing food, and for egg laying and larval

development. The queen termite resides in a royal

chamber, built to surround her pulsating, egg-producing
abdomen. Worker termites tend to her, removing the
eggs as she lays them to nursery chambers, where they
develop. The queen termite secretes a pheromone from
her body, which stimulates the workers to build pillars,
arches, and eventually walls around her to create the
royal chamber. Pheromone deposits may also help
direct the construction of walls and passageways
within the termite mound.
Macrotermes species

a Up to 114 in (3 cm)
b Savanna in Africa and Southeast Asia.
c Worker termites are usually pale brown to white and

are blind and wingless. Reproductive forms are often

darker and have compound eyes and two pairs of wings.

Termite mounds have an ingenious, built-in
ventilation system: variations in pressure or
temperature cause air to circulate, cooling and
freshening the colony. Wind, passing over openings
on top of the mound, decreases the air pressure,
drawing clean air into openings at the mounds
base and through its passageways.
warm, stale
air exits
cool, fresh
air enters


Macrotermes termite mounds can reach heights of
2023 ft (67 m). In dry areas, vertical shafts are
excavated to reach the water table, which can be
as much as 150 ft (45 m) below the ground.
royal cell
living chambers
and nursery

Many termites cultivate fungal fruiting
bodies on which they and their
nymphs feed. The fungi thrive on the
cellulose contained in chewed plant
matter supplied by the termites.

Wax combs
Honey bee

Common wasps are social insects that build nests out of paper to
house their colonies. Only the queen wasps (the sole egg layers)
survive the winter to found a new colony in spring. They
select a suitable site for the nest, either underground in
an abandoned animal burrow or in a sheltered
location, such as a garden shed. There they
begin to construct a nest from chewed
wood either from nearby trees or wood.
The wood pulp dries into paper, a
remarkably strong yet light building
material. Once complete, the nest
typically has a number of
horizontal layers of cells that
open downward. These house
the eggs and developing larvae
of the colony. Surrounding these
brood cells are a series of spiral
chambers, which provide strength to the
nest as well as trapping air for insulation.

Honey bee colonies live in beehives formed from several layers

of honeycomb. The honeycombs cells are used to house the
colonys developing brood and to store food in the form of pollen
and honey. Worker bees build the comb out of wax, which is
secreted from glands in their abdomen and manipulated with their
mouthparts. Workers produce different cells according to their
intended purpose. Cells for the larvae of drones (whose only duty
is to mate with the queen) are larger than those for worker larvae;
and those that will contain future queens are even bigger, oval in
shape, and are arranged vertically rather than horizontally.

Vespa vulgaris
8 1 2 in (11.5 cm) (worker);

2 4 in (1.52 cm) (queen)

b Nests underground or in buildings near yards,

woodland, and meadows in Europe, Asia, North Africa,
and North America.
c Wasp with black and yellow markings, a distinct
waist between the thorax and abdomen, and two pairs
of wings. The sting is on the tip of the abdomen.

Apis mellifera

a 1 2 1 in (1.52.5 cm)
b Worldwide. Some subspecies are native

to Europe, Africa, Middle East, and Asia.

Introduced to North and South America.

c Hairy, yellow-brown body. In workers,

the ovipositor is modified to form a sting.

d 185, 42425, 463

A single queen starts by
constructing a stalk
from which the nest is
suspended. She then
begins to add cells, laying
a single egg into each one.
When fully grown, the
workers complete the nest.

Common wasps use their jaws to
chew wood and place it in position
on the nest, then they use their
mouthparts to work the pulp to the
required thickness. The jaws of
wasps that use paper to build their
nests are shorter and broader than
those of wasps that build with mud.

Leaf nests
Weaver ants
Weaver ants, also called green or tree ants, use living leaves to
construct their nests in the tops of trees. The nests provide shelter
for the workers and developing young. A single colony of weaver
ants can consist of 100,000500,000 individuals and may have
up to 150 different nests in 12 or more trees. One of the colonys
nests will contain several egg-producing queen ants. This nest is
characterized by having more ant trails connecting with it than any
other. Workers travel along these trails to distribute eggs to other
nests. At the edge of the colonys territory there are barrack nests,
where older worker ants live and defend the boundary. Weaver
ants are voracious predators of pests that damage fruit crops,
such as citrus fruits, mangoes, and cashews. Fruit growers
therefore introduce weaver ant colonies into their plantations and
install bamboo bridges
between trees to help the ants
move around. The ants reduce
the need for chemical
When an ant finds two leaves with
sufficient flexibility to form a nest,
chains of its fellow worker ants line
up and straddle both leaves, pulling
their edges close. They hold them
there while other workers stick
the leaves together.


brush on
rear leg



As the colony grows, the workers gradually
demolish the innermost walls of the nest and build
new ones around the outside. A completed nest can
house 5,00010,000 wasps.

Honeycomb consists of
sheets of regular, sixsided cells. When a cell
is filled with mature
honey, or when a larva
is ready to pupate,
each individual cell is
capped with a wax lid.
Wild bee colonies
(right) live in hollow
trees or rock cavities.

Adult weaver ants use
sticky silk extruded
by the larvae to join
leaves together when
nest building. Unlike
other ant species,
the larvae do not
use their silk to
create cocoons.

Once several leaves have been glued into place, the
nest is complete. Worker ants move in with some of
the colonys eggs, which they tend through the
larval and pupa stages to adulthood.

Oecophylla species


The number of nests

that may be found in
a mature colony of weaver ants

a About 14 in (6 mm)
b Trees in tropical forests of Africa, Asia, and Australia.
c Relatively large, red-brown ants. Distinct separation

between head, thorax, and abdomen. Six legs attached to

the thoracic region. Antennae have 12 segments.


Paper houses
Common wasp


Second-hand homes
Shell-brooding cichlid
The shell-brooding cichlid uses empty snail shells as nurseries for
incubating its eggs. The female cichlids are attracted to nests of
shells collected by the much larger males. Larges males have an
advantage over smaller males in territorial disputes and when
competing for shells. Conversely, it is an advantage for females to
be small, so they can t deep inside the snail shells, where they lay
their eggs and remain for up to two weeks to brood them until the
hatchlings are ready to leave. There are also some medium-sized
males that will sneak into a resident males territory, where as
many as 14 females may be
Neolamprologus callipterus
nesting in different shells, in
a3 41 2 in (12 cm) (male);
an attempt to mate with the
1 4 in (4.5 cm) (female)
females. Another tactic is
b Fresh water of Lake Tanganyika,
employed by dwarf males,
central eastern Africa.
which are small enough to
c Gray-brown fish with elongated dorsal
enter a shell to fertilize the
fin. Males are much bigger than females
(sometimes up to 27 times larger).
females eggs.

Foam nursery
Gray foam-nest frog
Male cichlids collect empty
snail shells and carry them
back to a nesting area, which
they attempt to defend from
intruders. The largest males
tend to have more shells and
consequently attract more
females than smaller males.

The gray foam-nest frog breeds during a

brief window of time in the rainy season,
when seasonal pools are available. A
mating pair nds a location on a branch in
a tree overhanging a pool. There, the male
begins to create a nest by whipping uid
(a secretion produced by the female) into
a foamy substance with his long legs and
webbed feet. The pair may be joined by
single males that take part in the nest
building; in addition, these males release
sperm as the female deposits her eggs

Animal shelter
Gopher tortoise

Solid structure

The gopher tortoise uses its strong, stout

forelegs and wide, at claws to dig burrows,
about 15 ft (4.6 m) long and 61 2 ft (2 m)
deep, in sandy, well-drained soil. A tortoise
may have several active and inactive burrows
within its territory, retreating underground
to avoid predators and the midday sun as
well as to sleep at night and hibernate. A
total of 302 invertebrate and 60 vertebrate
species have been recorded as using
gopher tortoise burrows for shelter,
including frogs, lizards, snakes, and rats.

Also known as the hammerhead, the

hamerkop builds the largest roofed nest of
any bird. The nest is built in a fork between
tree branches. First, a platform is made
out of sticks, and then the sides are built
up into a deep basin. Next, a domed roof
of sticks and mud is added. The small
entrance is usually located at the side near
the base, as a precaution against predators.
Both the entrance tunnel and the nest
chamber are lined with mud. The nest
can reach 61 2 ft (2 m) high and wide and
may take up to six weeks to build. When
complete, it can support the weight of a
grown man on its roof. The whole structure
is decorated with unusual objects such as

into the foam. On average, the nests

of the gray foam-nest frog contain
approximately 850 eggs, which
take around 31 2 days to hatch into
tadpoles at a temperature of 77 F (25 C).
The tadpoles live in the foam for another
two days before emerging simultaneously
and dropping into the pool below.

Chiromantis xerampelina

a1 1314 3 in (4.57.5 cm) (male);

2 4 3 2 in (69 cm) (female)

b Subtropical and tropical forest, savanna, shrubland,

and grassland in central and southern East Africa.

c Gray-brown mottled frog with large eyes.

feathers, snakeskin, bones, and even manmade items. A pair of hamerkops may
build several nests within their territory, but
use only one. Other birds, such as eagle
owls and Egyptian geese, often move into
the spare nests.
Scopus umbretta

a 181 2 22 in (4756 cm)

b Freshwater wetland habitats in sub-Saharan Africa,

Madagascar, and southwestern Arabian Peninsula.

c Medium-sized wading bird related to pelicans. Brown

in color with a strong bill and large, shaggy crest at the
back of the head.

Hamerkops mate for life, and a pair labors for about
fours hours a day to build their massive nest. When it
is finished, the female lays between three and nine
brown eggs in the nest chamber.

Sandbank burrows
Carmine bee-eater

Gopherus polyphemus

a 10 in (25 cm)
b Sandy ridges, sand dunes, and longleaf pine forests

in the southeastern US.

c Tortoise with a brown-gray upper shell, dull yellow-

tan undershell, and gray-brown soft parts.

The female lays her
eggs into the foam
whipped up by the
males (top). The outer
layer of the foam
hardens into a nest
around a moist interior.
Oxygen diffuses
through the foam,
enabling the tadpoles
to breathe until they
drop into a pool of
water (above).

Carmine bee-eaters excavate horizontal

burrows in vertical sandbanks along rivers.
Their colonies may contain thousands of
individuals, often divided into smaller
groups known as clans. When starting
to excavate a nest burrow, a bird ies
repeatedly at the bank beak rst, until it
makes a slight depression onto which
it can cling. It then uses its bill and feet to
dig a narrow tunnel measuring 31 4 61 2 ft

(12 m) in length. Two of the carmine beeeaters toes are fused at the base, making
them perfect for using as a shovel.
Immature birds without a mate may help
their parents dig a new tunnel.

Merops nubicus

a 101 2 in (27 cm)

b Lowland river valleys, floodplains, and riverine forests

in sub-equatorial Africa.

c Colorful bird with a bright orange-red back, pink-red

chest, blue-green crown, and blue undertail. The
southern subspecies has a pink-red throat, the northern
subspecies a blue-green throat.

The white stork builds one of the largest

nests of all bird species. A pair mates for
life and tends to return to the same nest
site every year, adding a new layer of sticks
and earth to the huge nesting platform that
was used the previous year. Both males
and females take part in building the nest,
although the male collects most of the
materials. In addition to sticks, he may
select rags and paper discarded by
humans. Birds that remain faithful to their
nest sites tend to have a lower risk of
breeding failure than pairs that start anew

elsewhere. White storks also help boost

the survival of their eggs and chicks by
bringing fresh cattle dung to the nest to
help keep it warm.
Ciconia ciconia

a 31 4 4 ft (11.2 m)
b Breeds in Europe, northwest Africa, and southwest

Asia; winters in tropical to southern Africa and the Indian

subcontinent. Inhabits open farmland close to marshy
wetland feeding areas.

c Large wading bird. Predominantly white with black

flight feathers on its wings and a red bill and legs.
At Los Berruecos in Spains Extremadura region,
white storks nest out of harms way on the top of
dramatic granite boulders.


White storks often build their nests on the
roofs of buildings, such as churches or
towers. Because of their close association
with human habitation, storks have long
been considered a symbol of good luck and
are also the fabled deliverers of new babies
to people. In many parts of Europe, people
construct lofty platforms to encourage the
birds to nest. Nests on buildings tend to last
longer than those in trees because buildings
can better withstand the nests weight; one
nest taken down from a cathedral weighed
more than three-quarters of a ton.

Tree house
Pileated woodpecker
The pileated woodpecker uses its strong bill to excavate large
nest cavities in trees, typically 1434 80 ft (4.524 m) above the
ground. It may excavate a new nest each year or it may reuse
an existing hole for several years. The trees used for nests
vary according to region, but include aspen, western larch,
ponderosa pine, and Douglas r. They are often dead and
already hollow, making them easier to drill into than living, solid
trees. The nests frequently face east or south, to benet from
the warmth of the sun. Other birds, such as American kestrels,
wood ducks, and screech owls, may use old pileated
woodpecker holes for nesting once they have been vacated.
Pileated woodpeckers also chip away at trees to uncover food
in the form of carpenter ants and beetle larvae that live in the
wood. They pick these up with their sticky, barbed tongues.
Woodpeckers feet are adapted for walking up vertical tree
trunks by having two toes that point backward.
Both sexes assist with
building the nest cavity,
which is lined with wood
chippings. Between three
and five eggs are laid and
incubated alternately by
each parent.


High-rise living
White stork

Dryocopus pileatus

a 161 2 in (42 cm)

b Coniferous and deciduous forests of North

America, particularly southern Canada and the

eastern US.

c Large woodpecker with predominantly black

body and brilliant red head crest. Males have a red
moustache extending backward from the bill.


long tongue
wraps around
spongy skull

inner eyelid
tail braces
against tree
strong neck

Woodpeckers hammer trees 20 times a second at a speed of 15 mph

(24 kph). They have several adaptations to allow their bodies to deal
with the shock of hitting such a hard substance so fast and furiously:
the chisel-shaped bill is anchored to the skull by a thick bone to
prevent jarring; the skull is relatively thick and formed of spongy bone
that acts as a cushion for the brain; the muscles behind their bills and
in their necks contract just prior to the impact to help absorb the
pounding and transmit the shock through the whole body to protect
the head; and a third inner eyelid closes to restrain the eyeball and
shield the eye from ying splinters.


A male southern masked weaver displays by flapping
his wings beneath his finished nest (bottom left). A
rival male has only just started his nest (center), but
he may yet be successful, because females prefer
fresh green nests to older brown ones.

The southern masked weaver bends,

knots, and stitches pliable grasses into
a globe-shaped nest suspended from a
tree branch. Groups of male southern
masked weavers build their nests in the
same tree, selecting positions at the ends
of twigs as protection from tree-climbing
predators, such as snakes. Once the initial
hoop of woven grass is in place, the weaver
starts to construct the nesting chamber.
The length of the birds reach determines
the size of the spherical chamber, and a
porchlike entrance is added once the
nesting chamber is complete. When a
female moves in, she lines the nest with
soft grasses and feathers before laying
a clutch of about ve eggs.
Male weavers prefer to select fresh
green vegetation for their nests. Young

>>01 The male southern masked weaver
begins making its nest by selecting a thin
branch from which the nest will hang. >>02
Next, it constructs a vertical ring or hoop of
woven grass strips. >>03 It continues to add
to the hoop by winding and knotting more
strips with its bill.


leaves are more exible than older ones

and can be woven more easily. Also, the
females prefer nests made from newer
materials. Weavers use a complex array
of fastenings, including spiral binding and
knots such as the half hitch, overhand, and
slipknot. Birds that are experienced in
handling fresh green building materials are
better able to construct their woven nests
than those without similar experience.
Male weaver birds can y a total distance
of 230 miles (370 km) to collect materials
when building their nests.
Nest weaving has evolved independently
at least twice in birdsonce among the
weaver birds of Europe, Asia, and Africa,
and a second time among the orioles,
oropendolas, and caciques (both are New
World blackbirds) of North and South
America. In the case of the weaver birds,
it is the males that construct the nests,
whereas the female oropendolas and
caciques do most of the nest building.



Ploceus velatus

a 41 2 51 2 in (1114 cm)
b Southern African shrubland, savanna, grassland,

open woodland, inland wetland, and semi-desert.

c Female is greenish yellow; male in breeding coloration

has a black face, throat, and beak, a bright yellow crown
and chest, a greenish yellow back, and a red eye.

miles The total distance traveled

by male weaver birds to collect
fresh plant material when nest building.

There are around 60 species of
weaver birds in the genus Ploceus
and all share similar traits. To
collect nest material, male weavers,
for instance the village weaver
(P. cucullatus, above), land near
the base of a leaf, cut into it, and
fly off with a long, thin strip.
Female weavers, such as the
little weaver (P. luteolus, right),
inspect a potential mates nest
before deciding whether to
stay and breed.


Woven globes
Southern masked weaver


Decorated stage
Vogelkop bowerbird

Expandable home
Long-tailed tit

Male Vogelkop bowerbirds build elaborate display stages out of

sticks in order to attract mates. Their constructions are large and
impressive, built on the ground, and measuring up to 6 ft (1.8 m)
across and 4 ft (1.2 m) high. Populations of Vogelkop bowerbirds
differ in their bower design and taste for decoration. For example,
in the Arfak mountains, they tend to build huts with arched
entrances, which they adorn with brightly colored owers and
fruits; while in the Fakfak
Amblyornis inornata
mountains, they construct tall
a 10 in (25 cm)
spires or maypoles and
b Mountain habitats of the Vogelkop
decorate them with relatively
Peninsula of Irian Jaya, Indonesia.
drab decorations, such as snail
c Plain, olive-brown bird.
shells, nuts, and fungi.

The long-tailed tit builds a globe-shaped

nest out of spiders webs, mosses,
lichens, and feathers. Usually placed
in shrubs, such as gorse, bramble,
blackthorn, or hawthorn, the nest is held
together by the small, rough leaves of the
mosses, which hook onto the loops of the
stretchy spider silk. As the brood grows,
the nest expands to accommodate them.
The outside of the nest is covered with
silvery lichens and white spider cocoons,
both of which provide camouage by
reecting light off the exterior. The nest
is lined inside with a soft, warm layer of
2,000 or more feathers, which can
comprise about 40 percent of the nests
mass. Long-tailed tits adjust the thickness
of the insulating feather layer in their nest
according to prevailing environmental
conditions, the most important being
ambient temperature.
Aegithalos caudatus
Between 6 and 12 eggs are laid in the cosy nest. Once
the chicks have hatched, both parents work hard to
provide their growing brood with insects. Relatives, whose
own breeding attempts have failed, sometimes help.

Mud nest
House martin
House martins build a deep, cup-shaped
nest of mud underneath the eaves of
buildings. Successive layers of mud are
built up and the nest is lined with soft
materials, including grass and hair. A
narrow entrance hole at the top allows the
house martins in but prevents other larger
birds, such as sparrows, from entering
and occupying the nest. Relatives of the
house martin, such as other martins and
swallows, show an interesting spectrum
of nest-building activities. Their ancestors
were burrowers, a behavior that is still

a 56 in (1315 cm)
b Deciduous woodland, hedgerows, and scrub in

Europe and Asia.

c Small songbird, black and brown above, off-white

below, with a white crown on the top of the head.

followed by bank swallows. Others, such

as the purple martin and tree swallow,
make use of existing cavities. Barn and
cliff swallows both construct nests from
mud like the house martin, but the barn
swallows nest is a relatively shallow cup
perched on top of a beam, while the cliff
swallow sculpts a spherical nest with a
projecting entrance tunnel.
Delichon urbica

a 5 in (13 cm)
b Open habitats in Europe, North Africa, and temperate

Asia in the Northern Hemisphere summer. Overwinters in

similar habitats in sub-Saharan Africa and tropical Asia.

c Blue-black head, back, and tail, and a white rump

and underparts. Shallow fork to the tail.


House martins gather pellets of mud in their bills from
nearby ponds, streams, and puddles before adding
them to the nest. Both sexes participate in nest
building, and several breeding pairs may construct
their nests close together.

Thatched roof
Sociable weaver
Sociable weavers live communally in groups of up to 500 birds in
the avian equivalent of an apartment block. The huge nests
resemble haystacks placed in the branches of acacia trees. The
birds work to maintain their nests year-round because since the
thick, thatched roofs provide protection from the cold winter
nights and hot summer days that are typical of the desert. There
may be up to 300 openings in the underside of the nest, each
leading to a tunnel that terminates in a chamber. Sociable weavers
share these chambers with other birds, such as the South African
pygmy falcon, chats, nches,
Philetairus socius
and lovebirds, while vultures,
owls, and eagles may build
a 51 2 in (14 cm)
their nests on the roof.
b Open savanna grassland and thorn
scrubland in northern South Africa and
southern Namibia.

c A buff-brown bird with buff-white

underparts and a black chin.

Weavers use various materials when
building their nests. The initial
structure consists of large twigs and
stems, into which grasses are poked;
sharp spikes of straw deter predators
from the entrances, and the chambers
are lined with soft fur and cotton.

North American beavers are masters at

controlling water through construction. In
order to have sufciently deep ponds in
which to build their homes, known as
lodges, they rst build dams to control the
ow of water and raise its level. The largest
beaver dam so far discovered was at Three
Forks in Montana; it measured 2,130 ft
(650 m) long, 13 ft (4 m) high, and was
23 ft (7 m) thick at the base. In addition to
dams, beavers construct canals from
favored feeding
areas, along which
they oat food back
to the lodge.
Beavers can have a
positive effect on

the ecology and movement of water in their

environment. Their dams can help control
soil erosion and ooding, while their ponds
provide valuable wetland habitats for other
wildlife. However, their activities can also
be harmful. They may cause ooding, for
example if they block drains beneath
roads, and there may also be economic
damage to tree crops.
The ponds and lodges help protect
beavers against predators such as wolves,
lynx, and bears. They also provide shelter
from the cold in winter. When the ponds
freeze over, the beavers remain in their
lodges or move freely under the ice.

Beavers have strong skulls and large teeth, perfect
for cutting through tree trunks and eating the bark
and cambium (the soft layer immediately beneath
the bark). Their teeth continue growing throughout
their lives, but are worn down by gnawing.
Castor canadensis

a 3546 in (90117 cm)

b Wetland, streams, and rivers in North America,

except far north of Canada and parts of southern US

and Mexico.

c Large rodent with a waterproof brown-black coat,

short, rounded ears, and a broad, flat, scaly tail.
A beaver lodge may be located on an island behind
the dam, on the edge of the pond, or on the shore
of a lake. Some beavers excavate burrows rather
roof of lodge covered
than build a lodge.

beaver packs mud

to seal dam


The shape of a beaver
dam is determined by the
speed of water flow. In
slow water the dam is
straight, but in faster
rivers it is curved in order
to make it stronger.

Beavers are quick to rebuild their dams if they
become damaged. They drag tree trunks by their
teeth and carry mud and stones in their forepaws.

heavy stones
brace dam

sleeping chamber
above water, where
mother beaver
nurses her kits

with piles of sticks and

sealed with mud

underwater passage
food cache of
stored wood for
winter eating


Hydro engineering
North American beaver


House of straw
Eurasian harvest mouse
Pregnant female Eurasian harvest mice
weave spherical nests in which to give
birth and raise their young. The nests are
typically built 1251 in (30130 cm) above
the ground, attached to the stalks of reeds,
cereal crops, and grasses. The mouse
usually builds at nightrst, she shreds
grass leaves with her teeth, then she
weaves the strands into a hollow ball that
measures approximately 4 in (10 cm) in
diameter. More grasses are threaded into
the structure to build up several layers, and
the nished nest is lined with a soft bed of
shredded leaves and grass. The nest may
have several entrances, but the female
keeps these closed throughout the rst
week after she gives birth. Litters usually
consist of ve or six young.

Their mounds afford the
prairie dogs a good view
of the surrounding prairie.
They maintain a lookout
for potential predators,
which may include
coyotes, bobcats, eagles,
and hawks. If they spot
a predator approaching,
they give an alarm call
and dive underground.

Micromys minutus

a 2314 in (58 cm)

f Reed beds, hay meadows, cereal crops, and grassy

hedgerows in Europe (except most of Scandinavia) and

northern Asia.

c Small mouse with a blunt nose, short, rounded ears,

and a prehensile tail. Golden reddish brown fur with a

white underside.

a 1417 in (3543 cm)

f Dry, open, short- to mid-grass prairies from central
Texas to southern Canada.

c Rodent with brownish red coat and white underbelly.

Tail has a black tip. Body hairs have white tips in summer
and black tips in winter. Small eyes and ears.

Night nest

heavily grazed

grass roots bind surface

layers together

Colonies of black-tailed prairie dogs live in extensive underground

burrow systems called towns. Towns are divided into wards, and
the wards into coteries. Each coterie is home to several closely
related females, an unrelated adult male, and their offspring. A
coterie may have as many as 50 or 60 entrances into the burrow
system. The burrows are on average 161 2 33 ft (510 m) long and
61 2 934 ft (23 m) beneath the ground. Some entrances are
surrounded by a mound of earth 314 ft (1 m) high, resembling the
rim of a volcanic crater. The mound helps ventilate the burrow
system by altering the air ow over it, causing fresh air to be drawn
through the tunnels. Its steep sides also help protect the animals
from predators and ash ooding. Other entrances have a
shallower, domed mound, and some have no mound at all.
Cynomys ludovicianus

Most nights, chimpanzees build a new

nest in which to sleep. Their beds are
usually high up in trees, 3366 ft (1020 m)
off the ground, safely out of the reach of
predators, with females nesting higher
than males. They use tree branches and
leaves to construct a round platform
measuring between 2431 in (6080 cm)
in diameter. Nest building can take as little
as one to ve minutes, depending on the
experience of the builder. Chimpanzees
generally sleep for about 12 hours each
night and also rest in nests during the day,
between bouts of foraging. In addition,
they may retreat to bed if they are ill or
injured. Infants sleep with their mothers
until a new sibling arrives, at which point
they start to build their own nests, often
practicing their technique on the ground.
Biologists can use the number of fresh
nests in trees to estimate the number of
chimpanzees in an area, while the noise of
construction can also reveal their location.
Gorillas and orangutans construct arboreal
nests similar to those of chimpanzees,
although gorillas also nest on the ground.

entrance at top of mound

of excavated material

Underground town
Black-tailed prairie dog


passing place
in vertical
access tunnel

400 million

The largest
number of animals ever found
in a single prairie dog town,
in Texas.

hay in main

Black-tailed prairie dogs
spend time underground
in their burrow systems
at night, in the heat of
the day in summer,
and during inclement
weather in winter.


First, the chimpanzee pulls several thick branches together and
presses them down to make a stable platform. It then weaves
thinner branches and twigs around the edge and uses broken
twigs and leaves to provide padding for the center of the bed.

Pan troglodytes

a 2937 in (7395 cm)

f Gallery forest, rain forest, and woodland savanna

across equatorial Africa.

c Ape with black hair, sometimes tinged with gray or

brown. Infants have white tail tuft and face which
darkens with age.
d 247, 454, 460, 465, 480481
The daytime nests of chimpanzees may be
old rather than freshly built and are frequently in the
same tree in which the animals have been feeding.
They are often lower down than nighttime nests.

The Honduran white bat lives in rain forests

containing plenty of Heliconia plants, and it
uses their long, broad leaves to make tents
under which it roosts. The bat nibbles
through the veins on either side of the leaf
midrib, causing the leaf to fold down to
form an inverted V-shape. Each tent may
be inhabited by between one
and 12 bats, usually a single
male and his harem of females.
A colony of bats may have
several tents scattered
throughout its area of forest.
The bats use the tents during

the day when sleeping to provide

protection from the elements, such as
the sun and rain, and from predators,
which include opossums and snakes.
The green light ltering through the leaf
onto the white fur of the bat makes them
almost indistinguishable from their shelter.
They are so condent in their camouage
that the bats will take ight only if they
detect movement on the main stem of
their Heliconia plant.

Ectophylla alba

a 11 2 2 in (45 cm)
f Rain forest in lowland Central

America, including parts of Honduras,

Nicaragua, Costa Rica, and Panama.

c Small white bat with black wing

membranes and yellow-orange facial

features, including a triangular nose-leaf.

European mole
European moles spend most of their lives
underground. They are able to dig about
66 ft (20 m) of tunnels each day, throwing
up characteristic mounds of discarded soil
as they excavate, much to the chagrin of
gardeners attempting to achieve the
perfect lawn. A single moles tunnel
system may cover an area up to 1 3 mile
(0.5 km) across. They are solitary and
territorial, behaving aggressively to their
Talpa europaea

a 41 2 61 2 in (1116 cm)
f Rich, deep soils beneath arable fields, deciduous

woodland, and permanent pasture in temperate Europe.

c Cylindrical body covered with black velvety fur and

a bare nose with sensitive whiskers.


The moles front paws are shaped like

shovels, with ve strong claws. They are
permanently turned out in a perfect position
for pushing soil aside. Moles also have other
features that help them in their burrowing
lifestyletheir tiny eyes are almost
completely covered by fur, their nostrils
open to the side rather than the front, and
they have no external ears that might
otherwise get lled with earth.

neighbors and taking over their tunnels

if they leave or die. During the breeding
season, between March and May, male
moles may extend their burrows in search
of mates. Females give birth to between
two and seven young underground, where
they remain for about ve weeks before
striking out overland to establish their
own territories.
Moles immobilize earthworms that drop into their
tunnel systems by delivering a bite to the head.
Earthworms can sense the vibrations of a digging
mole and will endeavor to get out of the way.

Snowy lair
Ringed seal
All seals require access to the waters
surface in order to breathe. When
openings in the sea ice start to freeze over
in the fall, ringed seals create breathing
holes through the ice, using the claws of
their front ippers, so that they can pop up
for air. In the spring, females excavate lairs
or caves out of snow that has accumulated
in drifts above their breathing hole, a
behavior that is unique among seals. Here,
they give birth to a single pup and nurse it
for about 40 days. The lair helps the pup
survive the biting cold and offers some
protection from predators. However, polar
bears can detect the snow caves by their
scent and sometimes break through the
ceiling. An adult seal stands a chance of
escape by diving into the water through its
breathing hole, but if the pup is still too
young to swim it becomes easy prey.
Ringed seals often have more than one
lair, with adjacent lairs up to 234 miles
(4.5 km) apart.

stored in larder


A fresh molehill is a
sure sign that moles
are present in an area.
Beneath, a network of
shallow and deeper
tunnels leads to one or
more nest chambers filled
with dry plant matter.
This is where moles sleep
and where females raise
their young.

Ringed seals maintain breathing holes in a layer of
ice up to 61 2 ft (2 m) thick by digging with their
strong claws.
Phoca hispida

a 21 2 514 ft (0.81.6 m)
f Ice floes, landfast ice, and open water in the Arctic
Ocean, northern North Pacific, northern North Atlantic,
Baltic Sea, Bering Sea, and Sea of Okhotsk.

c Predominantly gray seal with whitish rings on its

back, a small head, and a short snout.

molehill of waste soil

above surface


Leafy tent
Honduran white bat



Feeding on









Grey herons typically smack large fishes against a
rock to kill them. They then flip them so that they
can be swallowed headfirst. This enables the heron
to eat the fish easily, without its fins, spines, and
scales sticking in the herons throat.



Feeding is a fundamental need of almost all animals because they require food to
provide energy for the various internal chemical processes essential for life. Even
though the details of how food is obtained and what food is eaten may vary
between animals, the basic processes involved are common to all of them. They
must locate food, capture or gather it, process it (by chewing, for example), ingest
it, and digest it. To achieve this, a range of feeding behaviors has evolved.

The starfish Pisaster
ochraceus feeds on
herbivores and so controls
biodiversity of rocky shores.


Food chains and webs

The physiological motivation to eatto replenish spent reserves, maintain

a positive energy balance, or fatten up for an event such as migration or
hibernationis controlled by a part of the brain called the hypothalamus
and by uctuating levels of hormones. But there are also other motivations
that drive animals to feed. Some may become conditioned to feed at a
particular time of day or in response to a particular sight, smell, or taste.
In some cases, simply coming across food is sufcient motivation to feed,
particularly if food is scarce. For many social species, seeing group mates
feeding is enough to stimulate feeding. In such situations, an animal may
eat more food than it would if
were it alone, and it is more
likely to eat a novel food if
others are eating it.

Feeding is essentially the transfer of energy from one organism, the

prey, to another, the predator. The energy in an ecosystem originates
from organisms that harness energy from the sun (plants) or from
chemical reactions (deep-sea bacteria). These primary producers are
eaten by primary consumers, which may in turn be eaten by other
consumers. In this way energy is
transferred through chains of
organisms. Each step in the chain
is called a trophic level. Typically,
the bottom of a chain (primary
producers) consists of a large
number of organisms, whereas
the top (apex predators) consists
of fewer, often larger, individuals.
Chains are components of more
complex food webs involving many
species and many levels.

Bears are more selective when food is
abundant, choosing to eat only the most
energy-rich parts of the fishes they catch.
Quality is less important than quantity when
food is scarce, and whole fishes are eaten.

Carnivores, herbivores, and omnivores

Animals can be classied according to their diet. Carnivores feed on other
animals. To do so they must be able to locate, kill, and eat their prey, and
so many have specialized prey-detection mechanisms, well-developed
weapons, and exhibit specialized feeding behaviors, such as stalking and
ambushing. Some carnivores feed on species far larger than themselves and
so must cooperate with one another to bring down prey. Herbivores feed on
plants, and many specialize in a particular part of the plant, such as fruits or
leaves. Like carnivores, herbivores require special feeding adaptations, such
as grinding teeth to chew tough plant material. Omnivores, scavengers, and
detritivores (which feed on
decaying organic matter)
are generalists; they eat a
wide variety of foodstuffs
and therefore often (but not
always) lack highly specialized
feeding adaptations. Instead,
they require behavioral and
anatomical exibility to locate,
process, and utilize a wide
variety of foods.
The dung of herbivores is an important
component of the diet of this Egyptian
vulture. The dung is rich in carotenoids,
which give the facial skin of the bird its
distinctive yellow color.

Relatively few very large blue whales (secondary
consumers) rely on billions of krill (primary
consumers), which in turn depend on a massive
amount of phytoplankton (primary producers).







baleen whales








detritus on
sea floor


The Antarctic Ocean food web may seem complex, but it is actually less
complicated than the food webs of many other ecosystems. Like all food
webs, it is delicately balanced and is vulnerable to disruption, from climate
change or overexploitation, for example. Disruption of one link in the web
might affect the entire web and could even lead to ecosystem collapse.

small toothed


When a kill is too large to eat in a single
meal, or if it is to dangerous to eat it where
it fell, solitary hunting carnivores like this
leopard will often store the carcass and
return to it later.

Competition and cooperation

Food resources are not innite, so individuals and populations have
to compete to secure their share. In some situations, one animal will be
dominant with respect to another and may be able to monopolize a food
source because of this heightened status. Or by defending a feeding
territory, an individual or group may be able to secure for itself all that it
needs. But when food is distributed across too large an area or numbers
of competitors are very high, territoriality may not be an effective strategy.
Species can coexist and minimize competition by each occupying a
particular niche in the environment. For example, they might feed on
slightly different foods or feed at different times or in different places.
















Detritivores, like wood lice,
feed on detritus such as
decaying wood, leaves, and
other plant material. In this
way they assist in the
recycling of plant material
back into the soil.

On the other hand,

individuals may need the
help of others of their
species to be successful.
Cooperation may increase
the chance that a territory
can be defended and also that food will be found, either because a group
can search a larger area or through direct communication about the location
of food. Cooperation may also allow animals to catch prey that they cannot
catch alone or allow a group to manage a resource for mutual benet.




Coyotes catch small mammals startled
by digging badgers and are more
successful when hunting with badgers
than when hunting alone.


An oak tree is a complex
habitat that provides various
niches for specialist
decomposers, herbivores,
omnivores, and carnivores.
These, in turn, are linked in
several food chains.

Foraging strategies
Carnivores and herbivores eat different foods but the basic principles of
feeding are common to all species. They need to nd food, process it, and
eat it. They need to make sure that they get the particular food they need
when they need it, and to do so they need to maximize the efciency of
their foraging. So just as a bear usually selects only the most nutritious
salmon, a goose usually grazes only the best-quality grass. Another
herbivore, the surgeon sh, is restricted to a low-quality diet of algae so it
simply eats a lot to obtain enough energy. Foraging strategies also need to
be exible, and animals need to learn what foods they can and cannot eat.


Some animals actively hunt for food; some
collect it from their surroundings; and some
remember previous locations of food







Some animals
avoid eating food
that has made
them ill in the past
or will not eat new
types of food they
have not eaten

Animals vary in
their feeding
behavior in
response to the
risk of predation or
of having food
stolen from them
by competitors

If food is abundant,
animals may share
it with others.
However, if food is
scarce, they may
eat a small amount
and cache the

Animals are
optimal foragers
and do not always
eat the first food
available if betterquality food can
be obtained easily
and safely


Foraging strategies are, in effect, a
series of decisions in which finding,
choosing, processing, and eating
involve a series of behavioral choices.


Optimal foragers
Animals are not indiscriminate in their feeding
behavior. Instead, they optimize efciency
through a trade-off between the costs of
nding and processing food, and the benet of
the energy gained from the food. For example,
small marine iguanas restrict themselves to
low-quality food found between the high- and
low-water lines because the extra energy they
would get if they fed on better-quality food
from beyond the low-water line is less than the
energy costs involved in diving to reach it.
Shore crabs show a similar behavior when
feeding on mussels (see below). Even sessile
fan worms feed optimally, balancing the energy
spent opening and closing their fans against
the amount of food in a current of water.
When feeding on
mussels, shore crabs
preferentially select
mussels 34 1 in
(22.5 cm) long
because they give the
highest energy profit,
taking into account their
calorific value (E)
and the time spent
opening them (T).

Ravens drop shells to open them.
They fly only as high as necessary
for the drop to break the shell.


A social motivation to feed



A physiological motivation to feed








Feeding on plants
Plant material is available as a potential food source in nearly all of the
environments of the world. Plants turn the energy of the sun into food
a process know as autotrophythat is usable by animals and underpins
almost all food chains. Therefore, the eating of plants is
fundamentally important to ecosystems.

Plants as food
Some animals specialize in eating one kind of plant,
or even one part of one kind of plant, while others
generalize, consuming a range of plants. Grazers eat
grassesand are often anatomically specialized for
thiswhile browsers consume plants selectively.
Granivores and frugivores eat seeds and fruits respectively,
and ultraspecialists like hummingbirds drink nectar from flowers. Nectar
is energy-rich, as are fruits, while seeds are rich in proteins and oils. Leaves
and grasses are not very nutritious, meaning animals often have to
consume large amounts of them. Plants are not passive in this relationship:
some deter predators with toxins or spines, while others encourage them
for the purposes of seeds pollination or dispersal, for example.

hummingbirds are hovering nectivores, with long beaks and

tongues for drinking nectar from flowers.

Furniture beetles are domestic pests. They burrow

into damp lumber, where their larvae eat the wood.


elephant skeleton
Elephants have large, ridged molariform teeth,
arranged on a conveyor; as the front teeth wear out,
those behind them slide forward, taking their place.

koalas are leaf eaters with a diet composed almost

entirely of eucalyptus leaves.

Wild mustangs are grazers, using their incisor teeth to clip the short
grasses that make up 8090 percent of their diet.


toxic leaves
As a defense against being eaten, the leaves of
many plant species are laced with toxic chemicals.
Some caterpillars that feed on these toxic leaves
disable the plants defenses by severing leaf veins
to slow the flow of the poison. Others excrete
the toxins they eat
and some, like the
Monarch butterfly
(left), use
them for
their own

types oF plant Food

When thinking of plants as food, there is a tendency
to picture animals grazing grasses. Plants, however,
are more than just green matter, and animals have
adapted to prey upon almost every part of the plant.
Insects, birds, and mammals consume nectar and
pollen, and a wide variety of animals eat flowers,
fruits, and seeds. Species such as pigs dig up plant
roots and tubers, and in desperation browsers will
chew twigs and bark. Termites and the larvae of
wood-boring beetles have evolved the ability to
digest cellulose, which allows them to eat wood.

gerenuks Feeding on acacia

Gerenuks are unique among browsing antelopes, as
they stand fully erect on their hind legs while leaning
against a tree with their front legs while feeding.
They can reach heights of more than 8 ft (2.5 m).

Some foods become available in a glut,

with availability far outstripping demand.
When the food is perishable, animals recruit
others to share the supply. If the food can
be stored, animals may benefit from
managing it. Some birds, for example,
store seeds and nuts in fall to prevent
winter food shortages. Other animals
manage quality rather than quantity.
Some grazers struggle to digest
grasses, but new grass is slightly
scattered pantry
During the fall months, a jay will store
more digestable, so some species
4,50011,000 acorns, each in a different site.
crop and rotate patches of grass,
Remarkably, it will retrieve them many months
later as food for its nestlings.
ensuring a fresh supply.

rotating harvest
In winter, brants move
as a flock from one
feeding field to the
next. They coordinate
their movements to
ensure that they do not
return to individual
fields too soon.

Brants wait for a period of 4 days before

they return to feed on land that they
have heavily grazed.

Mutual benefit
Some plants benefit from being food. Many flowers are pollinated by animals
moving between blooms as they eat nectar and pollen. Some seeds have a
tough coating that requires them to be digested before they will germinate,
and many seeds have far higher
germination rates if they have passed
through the gut of an animal. Also,
animals can carry the seeds a long
distance from the parent plant.
african elephant dung
Large seeds pass through an elephants gut and are
deposited to germinate in a pile of dung. Elephants
disperse the seeds of many tree species, including
some with no other known dispersal mechanism.

Plant defenses
While some plants do benefit from their relationships with herbivores, most
attempt to resist predatory animals. They defend themselves with toxins
and spines, or have a growth form that makes feeding difficult. Animals
can adapt to plant defensessome eat an antidote to plant toxins, some
develop immunity to them, and some utilize the poisons for defense. Some
herbivores have evolved to reach
past spines, some have harder teeth
to grind tough plants, and some, like
the chimpanzee, use tools to crack
open tough nuts.
The 20 in (50 cm) long tongue of the giraffe snakes
between thorns to pick small clumps of leaves.
Giraffe saliva is thick, coating spines eaten
accidentally so that they can be safely swallowed.

183 feeding on plants

Managing crops


Scouring ice
Antarctic sea urchin


Echinoderms such as sea urchins have an elaborate
apparatus for feeding called Aristotles Lantern. It is made
up of 50 skeletal elements that are largely internal, with
only the tips of the ve downward-pointing calcium
carbonate teeth, arranged in a downward-pointing cone,
protruding. In the center, a eshy substance serves as
a tongue. Sixty powerful muscles work together to push
the cone outward and to open and close the jaws.

All sea urchins are typically grazers. Crawling across the sea oor,
they use their powerful jaws, which consist of ve downwardpointing, chalky, bladelike teeth, to bite and scrape at food. The
Antarctic sea urchin scrapes food from the seabed and directly
from sea ice, feeding upon a lm of diatoms and on clumps
of red algae. Unlike other urchins, this species also consumes
the feces of Weddell seals. Urchins are able to use their
numerous tube feet to gather up material that has been stirred
up in the water.
Sterechinus neumayeri

a Up to 234 in (7 cm)
b Shallow waters of the Southern Ocean.
c Small, variable in color, ranging from greenish

gray to red or purpleviolet. Long spines extend from

its round shell.

Voracious swarming
Desert locust
During times of drought, the desert locust
is a solitary, drab, well-camouaged insect.
However, when the desert rains nally fall it
undergoes something of a transformation
in both appearance and behavior,
becoming both brightly colored and
highly gregarious. So dramatic is this
transformation that the two forms were
long thought to be distinct species. Rain
triggers voracious feeding among the
insects, causing them to lay down body

Groups of Antarctic sea
urchins congregate
at diatom mats, where
they graze on these
microscopic algae.

A swarm of locusts can cover up to 463 square
miles (1,200 square km) and will indiscriminately
consume leaves, stems, shoots, fruit, and seeds
of any plants in their path.
reserves and mate. Newly hatched
locusts do not y. These hoppers, as
they are known, form larger and larger
swarms, molting, growing, and eating
the equivalent of their own body mass
each day as they consume everything
green in their path.
Locusts are indiscriminate feeders,
eating leaves, stems, shoots, fruits, and
seeds of both noncrop and crop plants,
and they can cause devastation to the rural
economies of the countries they afict.
During their fth and nal molt, the
hoppers develop wings, and swarms of
tens of millions of individuals take to the
air, traveling up to 125 miles (200 km) per
day in search of fresh habitats in which
to continue to feed and breed. If this
coincides with a dry period, the emerging
nymphs will revert to the solitary form.
Locusta migratoria

a Up to 314 in (8 cm)
b Desert in North Africa.
c Winged insect with short antennae and variable

coloration depending upon life stage.

When grasshoppers crowd together, the
physical contact stimulates production
of pheromones, which in turn causes
the insects to swarm.

Probing proboscis
Cabbage white buttery

The proboscis of
this butterfly dips
repeatedly into the
multiple nectaries of
the flower, sipping
sweet nectar from
each of them.

Caterpillars of the cabbage white buttery are typical folivores (leaf

feeders). The female usually lays its eggs on the leaves of cabbages,
cauliower, or other similar plants. Once the eggs hatch, the green
caterpillars feed on the owers and leaves of their host plant. As a
free-ying buttery, the diet of the cabbage white changes, and it
then becomes a nectivore (nectar feeder).
Capable of powerful ight, the buttery searches for food plants
across a wide area and locates them by their scent. Having landed
on a suitable ower, it applies hydrostatic
pressure to two specially elongated mouthparts
Pieris brassicae
that are usually held coiled beneath the head.
a 221 2 in (5-6.5 cm) (wingspan)
Once straightened, these mouth parts zip
b Farmland, meadows, and parkland throughout
Europe, North Africa, and Asia to the Himalayas.
together to form a drinking tube, the proboscis,
which can be guided into the nectary of the
c Small, strong-flying butterfly, with black spots
on white forewings.
ower to extract the sweet nectar.

69 billion

The estimated
number of locusts in a swarm
that devastated parts of northwest
Africa in 2004.

This social insect uses sophisticated

communication to nd sources of food and
builds elaborate structures to process it.
Foraging worker bees visit up to 40 owers
per minute, collecting pollen and sucking
up nectar to be stored in the honey sack,
a specialized gut structure. Bees nd food
by sight and smell, but when a successful
bee returns to the hive it communicates to
other workers the location of nectar-rich
owers by means of the waggle dance (see
p.463). The way in which the dancing bee

moves communicates both the direction

and distance of the owers from the hive.
The colony is made up of sterile female
workers, male drones, and a queen. In the
hive, ower nectar is repeatedly swallowed
and regurgitated by the workers until it
becomes honey. Stored pollen and honey
are then fed by workers to the developing
larvae in their care. Some of the larvae are
put into separate chambers, and fed royal
jelly, a secretion from the feeding glands of
the workers themselves. These individuals
will develop to become future queens.
Apis mellifera

a 1 2 1 in (1.52.5 cm)
b Thought to have originated in India, now one of the

worlds most widespread domesticated insects.

c Small honey-brown-colored body, with sparse hairs

on its thorax and abdomen.
d 167, 42425, 463
Commercial beekeepers collect honey and
take it away from their hives. Hives may also
be moved between farms to maximize the
pollination of crops.


In recent years beekeepers have reported
the sudden disappearance of adult worker
bees from colonies that appear otherwise
to be doing well (they have plenty of
stored honey and a developing brood).
This phenomenon has been labeled
colony collapse disorder, or CCD. The
cause of CCD has yet to be determined, if
in fact there is a single cause. Suggested
causes include a resistant parasitic mite,
Varroa destructor, climate change, the use
of antibiotics, increased exposure to
pesticides, and viral, bacterial, and fungal
infections. However, the impact of CCD is
clear: this is a problem on a global scale
that is having a huge economic impact,
not just in lost revenues from honey
production but also because honey bees
pollinate so many important crop plants.

Bees collect pollen from the flowers they visit.
This is stored in the pollen basket, easily visible
as a yellow-orange ball on the hind legs. While
nectar is gathered as an energy source, pollen
is collected primarily for its protein and other
nutrients, and is used as food for larvae.

The hexagonal cells of a
honeycomb serve as both
a pantry in which to store
honey and pollen, and a
nursery in which the
young bees are raised.


The number
of bees in a
typical domestic beehive.


Processing pollen
Honey bee

Some honeypot ant workers (called repletes) have
enlarged abdomens filled with nectar, which they
regurgitate on demand for other members of the
colony. The repletes are fed by other workers, which
obtain food by gathering nectar from flowers, as well
as milking aphids and scale insects for honeydew
and preying on other invertebrates, such as termites.


Algae gardener
Blue damselsh

Reef grazing
Powder blue surgeonsh

The blue damselsh is a pugnacious herbivore that defends an

exclusive feeding territory. The impact of the territorial damselshes
upon their coral reef habitats is so great that they are often referred
to as being a keystone species. They feed selectively upon the
most nutritious parts of algae and sea grasses, which may in turn
benet from the shes habit of voiding their waste within the
territory. Through their selective grazing, damselshes inuence
the growth of their favorite food plants and control the growth
of algae; without this control, the algae would proliferate and
suppress the growth of the underlying coral.

Living on a diet of algae, which has limited nutritional value,

surgeonshes, or tangs, consume large volumes of food and have
an unusual range of gut symbionts (benecial organisms) to aid with
digestion. Surgeonshes also have beaklike mouths that are
equipped with a row of tiny, sharp teeth to scrape and bite at algal
turfs and mats. This beaklike mouth allows them to nip food items
from between obstacles, and their bite is so precise that they can
pick off algae from coral without damaging it. When food is scarce,
a surgeonsh will vigorously defend its territory, but if food is plentiful
they will occasionally feed in schools with others of their species,
and will even share a territory
with a blue damselsh (see left).
This strategy is advantageous
because the smaller damselsh
takes a relatively small share of
the food but is an aggressive
defender of the communal
patch. Competition between
surgeonshes species is
minimized because each
species in a community has
slightly different food
preferences. For example,
of the three species of
surgeonshes occurring
together on Florida coral reefs,
the blue tang prefers red algae,
the ocean surgeonshes prefer
green algae, and the
doctorshes specializes in
brown algae.


A blue damselfish tends its crop
of nutritious algae and defends it
against competitors. Without the
grazing action of damselfishes,
the coral reefs would become
overgrown with algae, which would
inhibit their growth.

Chrysiptera cyanea

a Up to 314 in (8.5 cm)

f Coral reefs of western Indo-Pacific,

to Micronesia and Samoa.

c Male is small and bright blue, with

yellow lips and tail. Females and juveniles
often have black spot at base of tail.

Acanthurus leucosternon

a Up to 10 in (25 cm)
f Coral reefs of the Indian Ocean

and Indonesia.

c Oval, powder-blue body with prominent

yellow dorsal fin, steep forehead, and black face.

When food is plentiful, surgeonfishes may
feed in schools. When food is scarce, they
are solitary feeders and each individual will
defend its territory against other surgeonfishes
and other species.

Arboreal browser
Monkey-tailed skink
The monkey-tailed skink is herbivorous and uses chemical cues
to identify edible leaves and owers. It spends the daylight hours
sheltering in the trunk of a hollow tree and emerges at night to
feed, using its prehensile tail to climb through vegetation. The tail
is crucial to the skinks arboreal lifestyle to such an extent that,
unlike most lizards, it is unable to shed it when attacked. Their
young are live born rather than hatched from an egg, and ride on
the back of their parent until they are able to climb themselves.

On the wetter islands, tortoises
migrate from one side to the
other to take advantage of fresh,
lush grass. They can also obtain
most of the moisture they need
from the dew on such vegetation.

Grazing giant
Galapagos tortoise
Galapagos tortoises graze on the leaves and fruits of more than
50 species of plants. They prefer lush grass, but their toothless
jaws are well adapted to cut and tear at all types of vegetable
matter, including tough bromeliads, fallen fruits, and the eshy
but prickly pads of cacti. On the wetter islands, the tortoises
drink water when it is
Geochelone nigra
available, but on the drier
a Up to 41 2 ft (1.4 m)
islands they obtain much of
f Galapagos Islands off the coast of
the liquid they need from their
Ecuador, South America.
food and can also store fat
c A giant brown tortoise.
within their large shells.

Diving for seaweed

Marine iguana
Marine iguanas are traditionally portrayed
as strong swimmers that dive to feed on
beds of offshore seaweeds. In fact, only
larger individualsthose above 4 lb (1.8
kg)habitually dive to feed. Smaller
individualsthose up to 234 lb (1.2 kg)
feed exclusively on the less lush seaweeds
of the intertidal areas, restricting their
feeding to the period of the day when the
tide is out and the sea is relatively calm.
The constraints of the tide present
difculties to small iguanas, who have to
limit their exposure to sea water so that their
body temperatures do not fall too farthe
sea around the Galapagos Islands is often
chilly, with a 5777 F (1425 C) annual
range. These animals must feed whenever
the tide is out, but if this is shortly after
dawn they risk being too cool and sluggish
to feed, and if the tide falls shortly before
dark, they may be chilled by splashing
waves and not have sufcient daylight
to reheat before nightfall. Small animals
huddle together at night to conserve heat.
Underwater foragers do not face
the same problems and typically bask
each morning before concentrating their
foraging time during the late morning and
around noon. This allows plenty of time to
bask and reheat during the early afternoon
when the sun still heats the island rocks.
Smaller iguanas do not feed underwater
because iguanas are actually relatively
poor swimmers, and smaller animals are
the poorest swimmers of all. They would
Using their claws and powerful legs,
large iguanas are able to resist the swell
of the sea to tear at submerged algal
mats and seaweed.

Corucia zebrata

a Up to 30 in (75 cm)
f Tropical rain forests on the
Solomon Islands.

c Long olive-green lizard with a

prehensile tail. Four limbs with sharp

claws and wedge-shaped head.

Monkey-tailed skinks use their
long, grasping tails and claws with
razor-sharp hooks to climb through
the tropical rain forest in search
of the edible leaves and flowers
on which they feed.

therefore take a long time to reach the

beds of seaweed or algae, risk being
dashed on the rocks by waves, and use
up a lot of their energy. Smaller bodies
cool faster than larger ones and therefore
small iguanas would be restricted to short
dives. So the limitations of their small size
seem to restrict their feeding opportunities
to the shore.
Amblyrhynchus cristatus

a 20 in31 4 ft (50100 cm)

f Galapagos Islands off the coast of South America.
c Black or dark gray-colored, large-headed lizard with

a scaly appearance and dorsal crest. The long whippy

tail is used for swimming.


Younger and smaller iguanas are unable to swim to
underwater seaweed mats and so must graze in the
intertidal area between the low- and high-water marks.

The number of minutes

that marine iguanas
can spend underwater while
diving for food.

By day, marine iguanas
bask on sun-warmed
rocks to raise their
body temperatures.
By night, they crowd
together in groups to
conserve heat.


On the drier Galapagos Islands
the cacti tend to be taller and
the tortoises have to rear up
and stretch their necks to feed.
The saddleback shape of the
shell allows greater neck
movement than the domed
shells found in tortoises that
live on wetter islands.


Larder tree
Acorn woodpecker
As its name would suggest, the acorn
woodpecker specializes in feeding on
the acorns of oak trees, although it
sometimes also feeds on insects and
the fruits of other plants. Like other
woodpeckers, it has a stiffened tail and
two forward-and two backward-pointing
toes on each foot to enable it to climb
the trunks of trees, and a specially
adapted bill and skull to allow it to drill
into wood.
In many parts of their range, extended
family groups of these birds may drill huge
numbers of holes into the trunk of a larder
tree, which they stuff full of acorns to serve
as a winter food store. In other areas, they
make use of natural crevices or even
suitable cracks and holes in manmade
structures. Groups of birds defend and
use this larder as a common resource and
may even breed communally, with one or
more pairs of birds in a group breeding
and being assisted by a number of
nonbreeding birds. Although they are often
sedentary birds, using the same larder for
a number of years, acorn woodpeckers
will move great distances if a failure of the
local acorn crop occurs.
Melanerpes formicivorus

a 71 2 9 in (1923 cm)
b Oak woodland in the southwest US and Central


c Black with white breast and cheek patches and red

crown and yellow throat.
arranging acorns
Each acorn woodpecker in a flock works to defend
the larder and to maintain the quality of the storage
holes. It will regularly rearrange their acorns into
smaller and smaller holes as they dry and shrink.


The number of acorns

often stored by acorn
woodpeckers in a
single tree.

Plucking fruits
Toco toucan
Although massive in appearance,
the prodigious bill of the toucan is
surprisingly fragile and lightweight.
It is composed of a hollow horny
sheath over crisscrossing
strengthening rods of bone,
and can easily be broken.
Toucans will forage for
food on the forest

floor, but they are essentially birds of the

forest canopy, where their long bill enables
them to reach out and
pluck fruits from hardto-reach branches.
A fruit held firmly in the
tip of the bill can be
maneuvered toward the throat
by the birds very long tongue, or
it may be flicked back with a toss of
the head and swallowed with a flourish.
Wild toucans eat the fruits of more than
100 species of forest plant and are almost
completely fruit eating, although they do
eat insects as a source of protein during
the breeding season.
Ramphastos toco

a 21231 2 in (5360 cm)

b Lowland rain forest (below 5,500 ft/1,700 m) in

northeast and central South America.

c Largest of the toucans, with a white breast and an

enormous yellow bill with a red keel and a large dark
spot at the end.

Hovering for nectar

Blue-chested hummingbird
The blue-chested hummingbird, like all
hummingbirds, has special adaptations to
its nectar-feeding lifestyle. The unique
arrangement of the wing bones and
joints enable these birds to
hover almost motionless,
and their extremely long bills
and even longer tongues
enable them to reach deep
into flowers to collect the
nectar that forms 90 percent
of their diet. However, nectar
lacks many essential
nutrients and hummingbirds
supplement their diets with
insects. But to spend all of
their time on the wing would
be too energy expensive, so
8590 percent of the time is
spent sitting still.

Amazilia amabilis

a 31 2 in (9 cm)
b Forested areas of Central America and northern

South America.

c Small, bronzy green bird with glittering green crown,

violet-blue chest, and long, pointed bill.

Although they occasionally eat insects,

owers, and sap, Bohemian waxwings are
almost entirely dependent on fruits, and
are one of very few temperate bird species
that can be described as frugivorous (fruiteating). Fruits tend to be abundant but
seasonal and patchily distributed. For this
reason, Bohemian waxwings are
gregarious throughout the year, often
breeding in loose colonies and forming
large and widely ranging ocks during the
nonbreeding season. Winter ocks of
Bohemian waxings can strip a tree bare of fruit
very quickly and travel significant distances
between trees. In this way they are important
in the dispersal of fruit seeds.

Secret store
Marsh tit
These avid food hoarders might hide up to
a hundred seeds in a single morning, and
thousands across a winter. Each seed is
stored in a different place, and seeds are
often some yards apart. Impressively, they
have the ability to relocate and retrieve
stored seeds days or weeks later. This
strategy allows them to take advantage
of food from sources that cannot be
consumed at a single sitting, such as a
sunower head or a backyard birdfeeder.
Not all tits have this ability; blue tits for

example do not store food and have poor

food-retrieval abilities. Comparisons of the
brains of storing and nonstoring species
have revealed that hoarders, such as
the marsh tit, have a well developed
hippocampal region, the area of the brain
associated with spatial memory.


waxwings often come into close contact

with humans, exploiting the fruits of
ornamental garden shrubs and trees.
Birds collect fruits while perched on a
branch, often plucking them and
swallowing them whole. When a bird
cannot reach berries from a perch, it will
briey hover to pluck them on the wing.
Most curiously, these social birds have
been observed to perch next to one
another along a branch and pass berries
backward down the line from one bird
to the next. Eating berries does, however,
have risks. Waxwings that eat fermented
fruits often succumb to intoxication,
which can be fatal.
Bombycilla garrulus

a 51 2 161 2 in (1417 cm)

b Forest in Europe, Asia, and North America.
c Gray-brown bird with distinctive crest and black

mask. Red wax spots on the secondary wing feathers.

Cone feeder
Red crossbill
Also known as the common crossbill, this
bird feeds almost exclusively on the seeds
of coniferous trees. The distinctive crossed
bill enables it to pry open the tough cones
of conifers before they ripen and open
naturally. This gives these birds an
advantage over other nches that share
their forest habitat but which do not have a
bill adapted to the task. Their resinous diet
means that crossbills must often visit water
to drink and clean their bills. By specializing
in this way, crossbills are able to breed
whenever the cone crop ripens, even
if this is in the middle of winter.

Garden birdfeeders save the lives of millions
of birds worldwide each year when there is
a shortage of wild food. Feeding in this way
can bring birds and humans closer together
than ever before. Provision of food all year
round has even changed the behavior of
some species, and
the list of regular
garden birds continues
to grow. However,
birdfeeders increase
the densities of birds
in a relatively small
area and therefore
may increase the
risk of disease
transmission, so good
feeder hygiene is
absolutely essential.

Loxia curvirostra

a 51 2 8 in (1420 cm)
b Coniferous forests in the Northern Hemisphere,

particularly Scandinavia and northern Britain.

c Large, stout finch, ranging in color from greenish

yellow to brick-red, with characteristically crossed,
pointed bill.

The marsh tit, like all of the tits, is an
accomplished acrobat and, if necessary, is
able to hang upside down to collect seeds.
After collecting the seeds, the marsh tit
hides each one in a different location, which
it remembers so that it can retrieve it later.
Parus palustris

a Up to 41 2 in (12 cm)
b Widespread in wooded areas of Europe and Asia.
c Small, agile bird with brown or beige upper parts,


A crossbill feeds by poking its closed bill into a
cone and then opening the bills crossed tips. This
pries open open the scale of the cone and enables
the bird to extract the seed.


Although crossed mandibles are a common
feature of the bills of all crossbills, the
particular size and shape of the bill varies longer upper bill
between species and even between
populations within a species. Each distinct
bill form is an evolutionary adaptation to
enable the birds to specialize in prying
apart the scales of their preferred type of
cone, whether it is spruce, pine, or larch.

buff-colored breast, and black cap.

Exclusive diet
Although they occasionally browse other tree species, koalas feed
almost exclusively on the brous leaves of eucalyptuses. Powerful
forelimbs, opposable digits, and strong claws enable them to move
with ease through trees to reach their food, which they nip off with
sharp incisors. Their modied cheek teeth, a single premolar, and
four molars with high crowns on each jaw, enable them to grind
the leaves to a smooth paste. Eucalyptus leaves are low in protein,
high in toxins, and difcult to digest, but the digestive system of the
koala is specially adapted to meet this challenge. The toxins are
deactivated and the paste is digested by bacterial fermentation in
a greatly enlarged cecum, which, at about 6 ft 6 in (2 m) long, is
the longest of any mammal.

After emerging from the pouch, a
young koala eats small amounts of its
mothers pap, a specialized form of
feces that is soft and runny. This
introduces into the youngs digestive
system the bacteria it will need to
digest eucalyptus itself. The pap is
also a rich source of protein for the
growing young koala.
Phascolarctos cinereus

a 2831 in (7278 cm)

b Eucalyptus forest and woodland in
eastern Australia.

c Small, gray bearlike marsupial with

characteristic pale ear tufts.


Gorging on berries
Bohemian waxwing


Flying frugivore
Great fruit-eating bat
Around one-third of the worlds bat species are
herbivores, feeding on fruits, leaves, pollen, and
nectar. The great fruit-eating bat is an important
pollinator and seed disperser for a wide range of
plant species, some of which are economically
important crops, for example mango, wild
banana, and durian. Flowers and fruits are
often locally superabundant, and vast ocks of
bats may descend upon a fruiting tree attracted
by one anothers calls and by the smell of the ripe
fruit. These ocks increase competition between
bats and attract predators, so many bats use their
teeth to tear off a fruit, which they carry away to
consume in solitary safety.
Artibeus lituratus

a 3 2 4 2 in (911 cm)
f Caves or large trees in forested areas of southern

Mexico through Central America, to southern Brazil, and

northern Argentina.

c Large brownish gray bat with a distinctive white

stripe above the eye.

Bamboo specialist
Giant panda
Classied as a bear, the panda may be
presumed to be a carnivore or an omnivore
rather than a herbivore, particularly given
that its diet is known to include meat,
shes, eggs, tubers, and plant material.
It also has bearlike teeth and a digestive
system more typical of a carnivore.
However, 99 percent of a pandas diet is
vegetarian and is composed of plants
of just one typebamboo. Having a
carnivores gut, the panda is not efcient at
digesting cellulose (plant glucose). It is able
to digest just 20 percent of the food it eats
while a more typical mammalian herbivore

Great fruiteating bats are
considered pests in
some areas because of
the devastating impact they
can have upon crops.

might digest 80 percent. The most proteinrich leaves of the plant are eaten
preferentially, and the bamboo is crushed
and ground by powerful muscled jaws
equipped with large at molars, but even
this processing does not make bamboo
a high-quality foodstuff, and the animal
needs to eat a lot of it. An adult panda can
consume up to 88 lbs (40 kg) of bamboo
per day and take 1214 hours doing it.
Stands of bamboo are synchronous in
their owering, death, and regeneration,
which means that pandas need to range
across relatively large areas of forest
supporting a range of bamboo species
to ensure a sufcient supply of food. They
feed mainly on the ground, but they are
also excellent climbers and able to swim.


At rst glance, the giant panda paw appears
to have an extra digit: ve ngers and one
thumb. This thumb, however, is actually an
elongated wrist bone, the radial sesamoid,
which is used by the panda as a pseudothumb. With it the panda is able to handle
food with considerable dexterity, gripping
bamboo shoots rmly in one hand, while
biting leaves from them.
true digits
have claws

Fruit and nectar eater

Kinkajous rarely leave the forest canopy,
and the bulk of their diet is forest fruit. They
are particularly fond of the sweeter, more
eshy fruits, which they collect in a manner
similar to monkeys; using dextrous hands
to process their food while hanging onto
the tree with their tail and feet. The muzzle
of a kinkajou is less pointed than that of its
close relative the racoon. This atter face
and extremely long tongue enables it to
lap nectar from owers. Kinkajous sleep
during the day, and feed at night.


Thumblike extension
of wrist bone without

Ailuropoda melanoleuca

a Up to 61/4 ft (1.9 m)
f Temperate forest with a dense bamboo

understory in southwest China.

c Long-haired, black and white bear.

The giant panda is so closely associated
with bamboo forests, and so reliant on
bamboo, that it is known to local people
as the bamboo bear.

Night grazer
Given their bulk, hippopotamuses eat
surprisingly little each day. Most of their
time is spent wallowing in warm water, but
they do not feed on aquatic plants. Each
night they leave the water and venture out
to feed almost exclusively on grasses,
which are digested in their multichambered
alimentary canal by the fermenting action
of microorganisms.

Remarkably, hippopotamuses consume

just 1.5 percent of their enormous body
mass each day, which would indicate that
their daytime wallowing and nocturnal
grazing is particularly energy-efcient.
Hippopotamus amphibius

a 1034 1114 ft (3.33.5 m)

f Rivers and lakes in West, Central, East, and South


c Heavy body, gray-brown above, and pink below, with

short legs. Eyes, ears, and nostrils are positioned at the
top of the massive head.

Potos flavus

a 31in5 ft (80115 cm)

f Tropical forests of Central and

northern South America.

c Mammal with dense golden fur,

and a superficial resemblance to both
a monkey and a cat.

Hippopotamuses have
formidable teeth, but
they use their thick lips
to nip at vegetation.
They have occasionally
been recorded
supplementing their
diet by scavenging on
the carcasses of
dead animals.

Chewing the cud

African buffalo
This buffalo has been described as one
of the most successful grazers in Africa.
It can be found in as diverse a range of
habitats as swamp, forest, and grassland.
It prefers taller grasses, using its long,
almost prehensile tongue to pull at clumps
of grasses that are then clipped by its wide
incisors and grazed to a pulp by its broad
molars. Through their grazing, herds of
buffaloes modify their habitat in a way that
makes it more suitable for other more
selective grazers, and this promotes the
growth of fresh food. As a result, herds of
buffaloes may be quite mobile. They are
able to avoid overheating by remaining
inactive by day and feeding at night.


The stomachs of ruminants (mammals that
chew the cud) have four chambers. Food
passes into the rumen and reticulum, where
digestion begins, fermentation takes place, and
solids are combined into a bolus. This bolus is
regurgitated and rechewed to break down the
plant material. This is then reswallowed and
passes into the third stomach, the omasum,
where liquid and essential minerals are
absorbed into the bloodstream. Remaining
solids pass into the nal stomach, the
abomasum, to be digested further.



Syncerus caffer

a 734 1114 ft (2.43.4 m)

f Grassland and woodland in sub-Saharan Africa.
c Thick-set, heavily built member of the cow family,


with a reddish brown coat and a pair of large upwardcurving horns with a large central boss.


One of the most successful of the grazing
vertebrates, the African buffalo prefers lush
grass but will graze coarse grass and herbs
when this is scarce.


passage of food
(first time)


passage of food
(second time)

Reaching the heights

The extraordinarily long neck of the giraffe
is an evolutionary adaptation that enables
it to compete with the many smaller
herbivores of the crowded savanna
ecosystem. Giraffes extreme height
enables them to reach above their
competitors and to selectively browse the
best parts of the tallest trees. They feed on
the fruits, owers, and fresh shoots of 40
60 different tree species, including mimosa,
commiphora (myrrh), and the spiny acacia.
Giraffes are able to nip leaves from
between the long thorns of acacia because
their long muzzles, exible lips, and long,
dextrous, tongues can reach deep
within clumps of tree branches and
because their lips and tongue are
protected from the thorns by thick,
horny bumps called papillae.
An adult giraffe consumes up to 77 lbs
(35 kg) of food each day, and to ensure
access to a sufcient quantity and quality,
roams widely. In lean times, giraffes eat
dried leaves, twigs, and even tree spines.
In common with other ruminants (see panel,
left), they rst chew and swallow their food,
then regurgitate and
rechew it several
times prior to
complete digestion.
Uniquely, they are
able to ruminate while
walking, an
adaptation which
perfectly suits their
nomadic lifestyle.
Giraffa camelopardalis

a 121 2 151 2 ft (3.84.7 m)

f Savanna, grassland, and open woodland in sub-

Saharan Africa.

c Large, long-legged and long-necked hoofed mammal

with a patchwork sandy-colored and brown coat.

Grazing the ocean floor

The only strictly marine mammalian herbivore, the
dugong, or sea cow, slowly meanders through the
sea-grass beds that are common in shallow, sandybottomed waters. As they rip out clumps of sea grass,
they leave behind trails of bare sand and uprooted
plants. This encourages regeneration of the sea
grass, promoting the nutritious new growth that they
favor. By repeatedly re-cropping the sea grass, they
maintain the quality of their food.


Thick, leathery lips and a very long tongue
enable the giraffe to selectively browse
tender leaves from around the sharp spines
of acacia trees.
Dugong dugon

a 814 13 ft (2.54 m)
f Shallow tropical waters of the

IndoPacific region.

c Long gray-green mammal with a

broad head and crescent-shaped tail.
Dugongs graze sandy-bottomed
waters and dig for rhizomes using
their flexible, thick upper lip.

Omnivores eat a wide variety of foodstuffs, often mixing

vegetable and animal material. they respond quickly to
changes in the availability of food and take advantage of
new food sources. their breadth of diet enables them to
survive in situations where specialists would struggle.

Feeding patterns

As generalists, omnivores tend to lack physical adaptations for feeding,

and their anatomy often combines that of carnivores and herbivores.
Omnivore teeth, a mixture of carnivore and herbivore teeth, are a good
example of this. Omnivores do exhibit behavioral adaptations, such as
an ability to manipulate and process a variety of foods, and they are often
good problem solversseveral omnivores
have learned to open trash cans. There are
drawbacks to having a relatively indiscriminate
diet. Animals feeding on seasonal foods often
suffer nutritional imbalances, but they are
physiologically adapted to withstand them.

Although generalist feeders may seem to be indiscriminate in their food

selection, most species do discriminate, choosing to eat less tough leaves,
for example. Filter-feeding mollusks ingest all small solids in their feeding
current, but have an internal mechanism for separating food from nonfood.
Many omnivores appear to
specialize at a particular point in
their annual cyclesome populations
of brown bear, for example, mainly
eat moths in the late summerso
variations are noticeable in omnivore
diets over long time periods.
all-day feeding
Chimpanzees feed all day, starting early and eating
almost everything in reach. As the day goes on,
they become more selective, choosing the ripest
fruits and most succulent leaves. Presumably, when
satisfied, they can afford to be more discerning.
adapting to a changing diet

hunting and feeding 194


Feeding behavior


plant material in diet


length of intestine


fear of the new

Rats are highly neophobic,
showing a strong reluctance to eat
unknown foods. This behavior
may have evolved to protect them
against accidental poisoning.

opportunism and adaptability

Although the most carnivorous bearand
dependant upon the fat of marine mammals for
much of its energythe polar bear is a highly
adaptable opportunist omnivore, and in
times of need will eat berries, kelp, and trash.
searching for food
Using its powerful and sensitive snout, the wild
boar searches inquisitively for buried food. It will dig
up roots and tubers, eat plants, kill small animals,
and even scavenge carcasses.


The gut length of starlings varies with diet.

Guts are longer in winter to permit a longer
digestion time for the plant material, which
is hard to digest, that forms the bulk of
their diet at this time.

filter feeding
Baleen whales, such as this Southern right
whale, feed by swimming open mouthed
through water containing zooplankton, and
other small pelagic animals. Water enters
the front of the mouth and then
passes out of the sides,
through baleen sieves.

Fan feeder
Peacock worm
The peacock worm lives in a muddy tube that extends from the
seabed. It constructs the tube itself, secreting mucus and binding
it with sand and mud. To feed, the worm comes to the mouth of
its tube, extending fans of nely divided featherlike tentacles
into the water. The worm waves its tentacles to trap material
suspended in the seawater, including grains of sediment
and plankton. It sorts its catch using the tiny hairlike
structures (cilia) that cover its tentacles. Edible particles
are transported toward the mouth to be eaten, while
larger, inedible particles are added to the tube that
surrounds the worm. If threatened, the worm will
quickly disappear into the safety of its tube.
Sabella pavonina

a Up to 12 in (30 cm)
f Mud and sand in shallow water

around the shores of Europe.

c Slender worm inhabiting muddy tube;

featherlike feeding appendages give it
flowerlike appearance.

By adding material
to their tubes as
they filter the sea
water around them,
peacock worms are
able to increase
their height.

Picking up a scent
Northern lobster

Predatory pollen lover

Soldier beetles

During its larval stage, the northern lobster, or American lobster,

is an opportunist carnivore, feeding on plankton. As it matures,
this crustacean becomes a generalist omnivore. It will scavenge
if necessary but prefers to hunt for its food, which includes
crabs, mollusks, bristleworms, echinoderms, and sometimes
other lobsters. Usually a nocturnal hunter, it locates its prey by
detecting scents in the water. The lobsters long antennae
and shorter antennules are incredibly sensitive and allow it to
discriminate between the odors of different prey. It has many
mouthparts, with differing functions, which include holding food
or passing it to the lobsters jaws, where food is crushed and
ingested. Thriving in cold, shallow waters, the northern
lobster is ercely territorial, occupying burrows and
crevices on the seaoor.

Both the gray-brown larvae and vividly

colored adult soldier beetles are voracious
carnivores. Adult females lay eggs in soil.
As they develop, the wormlike larvae
spend most of their time among leaf
litter, where they prey upon
invertebrates, including
snails and slugs.
Soldier beetles are
important predators of
caterpillars, aphids, and
other insects. Adult soldier
beetles congregate on the owers
of herbaceous plants, particularly
pollen and nectar-rich species such as
goldenrod and umbillifers, where they mate
and prey on other visiting insects. While
inhabiting the owers, the beetles become
omnivores, supplementing their diet with
pollen and nectar.
Soldier beetles are named for their
coloration, which resembles a military
uniform. The bright colors are a warning
to predators that the beetles taste
unpleasant. These insects have exible
wing covers, giving them their alternate
name of leatherwings, and a long body
that, unlike those of other beetles, is soft.

Homarus americanus

a 824 in (2060 cm)

f In crevices in ocean water along the

North Atlantic coasts of Canada and the US.

c Long blue and red crustacean with a

massive pair of front claws.


The northern lobster has two
massive front claws. One is used for
crushing and breaking the shells of
its prey, while the other has a sharp
cutting edge for tearing at flesh.

Family Cantharidae

a Up to 3 4 in (2 cm)
f Herbaceous vegetation in sunny positions worldwide.
c Slender, elongate beetles with parallel elytra

(forewings) and distinctive red, yellow, and black

coloration. Head has curved jaws and slender antennae.

Colorful soldier beetles are welcomed by
gardeners, since they are predators of
aphids and caterpillars, which are
perceived as pests.


This peacock worm has
extended its feathery,
banded tentacles into
the water to feed.

Lethal teeth
Piranhas have a reputation for being fearsome
predators. Their typically sharp, slicing teeth, which
interlock when their mouths are closed, and their
protruding lower jaws make them efcient biters.
Piranhas also have a keen sense of smell, which is
used to locate food even in the murky waters of
ooded rivers. They usually prey on shes and
invertebrates that are smaller than themselves,
but they occasionally feed in a frenzied pack,

and also sometimes kill and eat larger animals, such

as capybaras, horses, and even humans. However,
there is a less erce side to these sh. During
ooding, piranhas move out into inundated areas and
feed on decaying vegetation and the seeds and fruits
of forest plants. The pacus, those piranha species
that specialize to a greater extent on fruits and seeds,
tend to have atter, thicker teeth more suited to
crushing seeds than to biting esh.

Serrasalmus species

a Up to 13 in (33 cm)
b Rivers of South America east of the Andes.
c A somewhat round, laterally flattened silver fish with contrasting dark

Coral crunching
Bullethead parrotsh
Like all parrotshes, the bullethead parrotsh is best described as
an omnivorous herbivore. Parrotshes main food is marine algae,
which they graze from coral reefs, thereby preventing algae from
smothering the coral. However, parrotshes also eat living and
dead coral. They do this to gain access to micro-algae that
colonize dead coral and to the symbiotic zooxanthella (microalgae) of living coral. To enable them to bite hard coral, the
relatively small teeth of parrotshes are fused into beaklike plates.
These wear quickly and so grow constantly. To crush the coral they
have additional pharyngeal teeth in their throats. As a result of their
feeding, they are considered a major erosion
force in reef systems, but the ne coral sand
they excrete may help stabilize the reef.

fins and belly. Both jaws have a single row of sharp, interlocking teeth
used for puncturing and shearing.

A large mouth and interlocking,
razor-sharp teeth enable this
piranha to tear chunks of flesh
from larger animals and
even humans.

Chlorurus sordidus
The sharp biting teeth of piranhas are
perfect for stripping the flesh from
forest fruits. Some species have
thicker crushing teeth to break up
the seeds inside.

a Up to 16 in (40 cm)
b Shallow coral reefs of the Indo-

Pacific region.

c Males are blue with yellow and orange

markings and blue lips, Females are reddish
brown with red lips.

Parrotfish erode dead
coral by biting into it
with their beaklike
plates at rates of up to
78 bites per minute.

This giant sh feeds on both planktonic

plants and animals as well as smaller sh,
crustaceans, and squids. Whale sharks
migrate through the oceans of the world
to exploit areas of rich feeding such as
the Australian Ningaloo Reef. There,
they congregate to feast on the plankton
explosion associated with the mass
spawning of corals.
Whale sharks are thought to be able
to determine the best feeding areas by
the use of olfactory cues. They usually
feed by cruising slowly through food-rich
waters, passing huge volumes of water
in through their mouths and out through
their gills, where food particles are
trapped for swallowing. Whale sharks
have also been seen using their mouths
like a giant bucket, swimming upward
through a dense patch of food to engulf it.

Whale sharks are not the only
filter-feeding sharks. Basking
sharks (left) and megamouth
sharks (below) also have huge
mouths and enormous gill arches.
Such adaptations enable them
to be efficient filter feeders.

5 ft

The approximate
width of a whale sharks
mouth, through which it
sucks water containing food.


The huge, gaping mouth of a whale shark scoops
up a school of small fish. All the water the shark
swallows will be pushed out through the gills, while
the fish will be trapped by sievelike membranes.

Whale and basking sharks, two of the three lter-feeding sharks,

synchronize gulping in water and opening their gills. They also create
suction to draw in water by expanding their buccal cavity (throat), or
gulping air at the water surface. Water pumped through the gills passes
through a sieve of bony projections termed gill rakers, which trap any
potential food particles larger than 1 8 in (3 mm). Basking sharks shed
their gill rakers in winter and grow a new set each spring, suggesting
that they are seasonal feeders The third lter feeder, the megamouth
shark, has softer gill rakers and may use its silvery palate and a
luminescent stripe above its lips as a lure to prey.

Rhincodon typus

a 3966 ft (1220 m)
b Worldwide, in tropical and temperate waters.
c The worlds largest fish, having a huge head and

prominent ridges running along its gray and brown body

which is patterned with white spots.

Mixed diet
Brazilian tree frog
While the tadpoles of many species of frogs are
omnivorous, feeding on vegetable material, carrion,
invertebrates, and even cannibalizing one another,
adult frogs are almost
entirely carnivorous.
Fruit-eating frogs, only
However, the Brazilian tree
11 2 in (4 cm) long, are able
frog is the exception. It
to eat berries that are up to
2 in (1 cm) in diameter.
rests in the pools of water


As critically endangered species, all marine
turtles are listed by CITES (the Convention
on International Trade in Endangered
Species), and their trade, capture, and
harassment is banned under international
law. However, cessation of trade alone is
unlikely to prevent the extinction of the
hawksbill turtletheir nesting beaches (in
some 60 countries) must also be protected.
They are still illegally hunted for food and
eggs, and the intricate patterning of their
shell is particularly sought after for jewelery
and decorative furniture.

Toxin tolerance
Hawksbill turtle
Hawksbill turtles feed among the shallow
algal beds of inshore waters or around
coral reefs. They consume a wide variety
of foods, including seaweed, mollusks,
crustaceans, and small shes. Their
strong, sharp beaks are used to bite
off chunks of both soft and hard coral
colonies. When feeding on jellyshes,
such as the deadly Portuguese man-ofwar, turtles close their eyes as a defense
against being stung. The bulk of their diet,
however, consists of sponges, many of
which are highly toxic and packed with
siliceous structures, called spicules, that
are as sharp as shards of glass. Neither
defense appears to deter the turtle.
Eretmochelys imbricata

a 23ft 3 in (0.61 m)
b Shallow reefs and inshore areas of subtropical

and tropical waters worldwide.

c The smallest marine turtle, recognized by its

shell shape, with distinctive central ridge, serrated
edge, and overlapping scutes.

that form within bromeliads during the day and

forages by night, when it eats invertebrates, but its
diet also includes the brightly colored fruits and
seeds of a range of plants. The seeds of one of its
preferred plants, Anthurium harrisii (an arum), are
known to germinate when they have been defecated
by the frog, so it
Xenohyla truncata
is thought that
this might be an
a Up to 11 2 in (4 cm)
important method
b Subtropical and tropical marshes
of Brazil.
of dispersal for this
c Small brown tree frog.
particular plant.


Filter feeding
Whale shark


Changing tastes
Bearded dragon
These lizards alter their feeding habits during
their lives. Juvenile bearded dragons are largely
insectivorous so they are considered carnivores.
They hunt small insects on the ground and in trees,
constantly turning their heads as they look for prey
and, when they have spotted an insect, bursting into
activity to chase and catch it. Between periods of
hunting, the young lizards rest and bask, often in
trees. As the lizards grow, their diet changes and

they begin to consume more and more vegetable

material, including leaves, fruits, and owers. These
plant parts are bitten off by powerful jaws. Bearded
dragons have rounded rather than pointed snouts, a
typical shape among herbivorous lizards. The tongue
also plays an important part in their
feeding: it can be icked out to
taste a food to check if it is
edible, and it can also be used
to pick up fallen fruits and owers.

Ground feeding
Southern cassowary
Being ightless and too large to climb,
the southern cassowary feeds on
the ground, searching through the
leaf litter to nd fungi, snails,
insects, and small animals,
such as frogs, to eat. It
sifts through the litter by
scratching at it with its
feet and turning it over
with its bill, but it also uses the
calcied cartilage casque that projects
from the top of its head as a shovel.

Amphibolurus barbatus

a 1012 in (2530 cm)

f Desert, scrubland, and forest in

eastern Australia.

c Gray, brown, or black lizard with a

short tail, heavy body and head. Yellow

lining to the mouth and a beard of throat

Adult bearded dragons still consume
animal prey, but their larger size
allows them to diversify, feeding on
larger insects, like this one, smaller
reptiles, and small mammals.

Flexible feeder
African harrier-hawk
Palm nuts and other fruits are often
eaten by the African harrier-hawk, or
gymnogene, but it is also an accomplished
hunter, and includes insects, small
mammals such as rabbits, birds and their
eggs, bats, and reptiles in its varied diet.
A harrier-hawk will walk on the ground to
search out food, or undertake low foraging
ights over patches of short vegetation,
pouncing on its prey. However, most often
this species is observed clambering
through trees and shrubs to forage using
its wings, feet, and bill to steady itself as it
climbs. The hawks slender bill is used to
probe beneath bark and in crevices to nd
insect larvae and small reptiles. There are
also observations of the bird hanging
upside down from a gray-headed social
weavers nest, and grabbing an adult bird
in its bill as it exited the nest hole.
The most remarkable thing about the
African harrier-hawk is its double-jointed
ankles; the joints bend both forward and
backward. The bird uses its long, exible
legs to reach at seemingly impossible
angles in search of hidden prey. Using
this technique, African harrier-hawks
have been seen hanging from the nests
of weaver birds and reaching deep inside
them to nd the eggs and young.

The combination of small head, long
neck, slender bill, and long, flexible
legs make the African harrier-hawk
skilled at finding food in small holes
and crevices.

Polyboroides typus

a 2426 in (6066 cm)

b Forest, woodland, and grassland in

sub-Saharan Africa.

c Medium-sized gray raptor with naked

yellow face and long yellow legs.

Male southern cassowaries generally have larger
casques than females, and the casques continue
to grow as the birds age. The red and blue
coloration of the skin on its neck indicates that
it is at least three years old.
Casuarius casuarius

a Up to 51 2 ft (1.7 m)
f Dense tropical forest in New Guinea and Australia.
c Large, thick-legged bird with solid crest or casque,

brown and black plumage, and striking red and blue skin
on the neck and face.

Soil supplements
Red and green macaw
In common with some other herbivores,
red and green macaws exhibit geophagy,
literally, soil eating. Large ocks of red
and green macaws (and other parrot
species) congregate on the eroded faces
of riverbanks and mud cliffs to socialize
and sometimes to nest in crevices, but
most commonly in order to eat the soil.
The macaw diet is composed largely
of fruits, seeds, berries, and nuts. Using
their keen eyesight to forage in the forest
canopy, groups of red and green macaws
locate food and dextrously pluck fruits
from the trees. Using their feet and sharp,
hooked bill they tear open the esh so that
they can get at the hard seed or nut within.
The macaws bill can generate an
enormous biting force, making it able to
crack even the hardest nuts.
In order to avoid competition with other
forest herbivores, for example monkeys
like the saki, macaws eat underripe fruits
and plants that are chemically defended
and generally unpalatable, or even toxic
to other animals. It is possible that
macaws eat soil to supplement the
mineral content of their diet, but it has
also been shown that by electing to eat
the particular soil type found at clay
licks, the birds are able to neutralize
the toxins in their gut.
Ara chloropterus

a Up to 3 ft 3 in (1 m)
f Tropical forest in eastern Central America

and northeastern South America.

c Large red, blue, and green parrot with

a naked white face striped with small
red feathers.
d 465


The pressure, in pounds

per square inch, that can
be generated by a macaws bill.

Fruit and meat


Seasonal food
Great bustard

Like all parrots, the kea eats fruits, nuts,

and seeds, but unlike other parrots, it
has a taste for meat. Some dig into the
nesting burrows of sooty shearwaters to
kill and eat their chicks. Others, living in
the hills and mountainssometimes
above the snow linescavenge the
carcasses of dead sheep. In the past,
keas were seen eating the fat from the
backs of live sheep, as a result of which
they were hunted almost to extinction.

Historically a bird of open grassland, the

great bustard is well adapted to survive
in the modern agricultural landscape,
particularly in areas of cereal growing.
However, its persecution by humans has
reduced its population considerably.
Young bustards are almost entirely
insectivorous, but as they age, the birds
include increasing amounts of vegetable
material in their diet. They forage in open
areas, meandering in loose ocks and
picking at selected food items. The birds

Nestor notabilis

a 18 in (46 cm)
f Wooded and alpine areas of New Zealands

Macaws regularly travel considerable distances to gather
at favored clay licks and eat the soil. The soil may detoxify
toxins from some of the plants in the macaws diet,
and soil eating may therefore be viewed as a form of

adapt their diets in response to

uctuations in food availability associated
with seasonal agricultural practices and
a habitat dominated by annual plants. In
summer they consume more insects
(particularly beetles and their larvae) and
also take small vertebrates, such as frogs
and mice; in spring, fall, and winter they
are almost exclusively vegetarian, with
a very varied plant diet, including
cereal crops.



Otis tarda

a Up to 31 2 ft (1.1 m)
f Scattered populations are found across the

grasslands of Europe and Asia.

c Robust reddish brown and gray bird. Breeding males

grow moustachial whiskers 8 in (20 cm) long.

South Island.

c Olive-green parrot with scarlet underwings.

The worlds heaviest flying birds,
male great bustards congregate
at leks (communal display sites)
and compete for the attention
of females.

The keas elongated bill is perfect for biting,
tearing, and lifting a variety of prey and objects,
even prying rubber parts from cars, a behavior that
has given it a reputation as a vandal.

After an absence of almost 200 years,

the great bustard might once again become
a familiar bird in parts of Britain. A program
of reintroduction using birds of Russian
origin (genetically the most similar to
historical UK populations) was started and
in 2007 a released bird laid eggs, although
unfortunately they did not hatch. Extensive
surveys determined that in the release
areas, insect numbers in the summer would
feed the growing chicks, but it may be
necessary to manage winter feeding areas
alongside crops to ensure the birds survival
over winter.


unfussy eater
Virginia opossum
The nocturnal Virginia opossum is
usually ground dwelling, but it is also an
accomplished climber. It has a prehensile
tail and very sharp claws on all of its toes
except the large toe of the hind foot, which
is opposable, like a thumb, and is therefore
extremely useful in grasping tree branches

while searching for food. With excellent

night vision but poor distance vision,
opossums are more likely to use touch and
smell to locate food. The Virginia opossum
eats a huge variety of food, from fruits,
seeds, and plant material, to eggs, insects,
and small vertebrates, including some
reptiles, amphibians, and other mammals.
It is even able to prey on poisonous
snakes, having a higher resistance to
snake venom than other mammals.

Shovel snout
White-nosed coati
A member of the racoon family, the whitenosed coati spends its days in a bustle of
gregarious activity as it searches, often in
groups, for food. It forages for its food,
mostly on the ground and usually by
digging and snuffling through the leaf litter,
although it is also a skilled climber.

Didelphis virginiana

a 15211 2 in (3855 cm)

b Forest, woodland, and farmland in

coatis eat nuts, fruit, carrion, eggs, and

small animals, such as insects and other
invertebrates and small reptiles, which they
sniff out with their excellent sense of smell
and highly developed snout. The snout is
shovel shaped and long, protruding
beyond the lower jaw. It is muscular and
flexible and can be pushed into crevices
or under bark in the search for food. coati
teeth differ from those of most other
mammalian carnivores, in that the molars
are flattened for crushing and grinding,
rather than for shearing flesh.
Nasua narica

Central America, the US, and southern

states of Canada.

a 21 2 41 4 ft (0.81.3 m)
b Wooded areas of southeast Arizona, US, and Mexico,

c Ratlike marsupial with silver-furred

face and naked, prehensile tail.

through Central America to Panama and northeast


c Long-nosed, long-tailed, slender carnivore with

a distinctive white muzzle and white face pattern.

Cleaning up Carrion
Virginia opossums are extreme
generalists and will eat almost
anything. They are also opportunists
and scavenge, taking advantage of
carrion whenever they find it.

fond of fruiT
The white-nosed coati can climb to reach fruits
and other arboreal foods, using its long tail
to help it balance; however, this species is
primarily a ground forager.

Tireless forager
Wild boar
To obtain the high levels of protein they
need to survive, wild boars are generalist
omnivores, using their snouts to locate
and then dig out buried food. Their ability
to find food in a wide range of habitats
explains their global success. Ranging
widely, and foraging from dawn to dusk,
wild boars rest during the middle of the
day and night. Plants, especially fruits and
nuts, constitute 90 percent of their diet.
The remaining 10 percent has been
known to include insects, eggs, small
vertebrates, carrion, and even refuse,
their exact diet varying with the seasons.


Three hundred years ago, wild boars in
Britain had been hunted to extinction, but
in the 1990s they were again found in the
wild. It is believed that some animals may
have escaped or been released from the
wild boar farms that were popular ventures
at the time. Significant numbers were then
found in many rural areas throughout
southern and western England. While some
view it as a welcome return, many farmers
disagree, citing damage to crops and
harassment of their cattle.

Sus scrofa

a 331 2 ft (0.91.1 m)
b Woodland in Europe, North Africa, and Asia.
c Typically porcine with large head and compact body,

brown and black in color and sparsely bristled.

d 485
an advenTurouS palaTe
Young wild boars are even less particular about
what they eat than their parents, which increases
their chance of survival in an environment where
food availability fluctuates.

Huge appetite
Blue whale
Believed to be the largest animal ever to
have lived, the blue whale is a filter feeder
that migrates to the southern oceans each
summer to feed mainly on krill (small
shrimps), and also consume other
crustaceans, squids, and small fishes
associated with krill swarms. During the
summer feeding season, a blue whale can
consume up to 4.4 tons (4 tonnes) of krill
every day. This allows the whales to lay
down reserves of fat and oil, which will
provide energy during the rest of the year
when food is less abundant. A feeding
whale opens its mouth, relaxes its throat,
and takes in a vast volume of water and
associated food. It then closes its mouth,
tightens its throat, and uses its massive

tongue to force the water out through

huge baleen sieves that hang from its
upper jaws. The krill are trapped against
the inner surface of the baleen sieves
and are then swallowed by the whale.
Blue whales can be found in oceans
all over the world, although the three
primary populations are in the North
Atlantic, North Pacific, and the Southern
Hemisphere. Typically living in groups
of two or three animals, they mate and
calve in tropical to temperate waters in the
winter months and feed in polar waters in
the summer.
Balaenoptera musculus

a 7989 ft (2427 m)
b May be found in all of the worlds oceans, although

absolute numbers are small.

c Mottled bluish gray; more slender in appearance than

other whales.

2,204 lb

The amount of food needed

to fill up a blue whales stomach.
geTTing a mouTHful
Krill undertake a daily
migration between deep
water and the ocean
surface. To reach them,
blue whales feed at
depths of 330 ft (100 m)
by day and close to the
surface at night.

Although they are often perceived as

fearsome carnivores, brown bears are
actually omnivores, with a diet that is at
times composed largely of plant material.
Their diet varies with the season and
geographical location, but tends to include
green vegetation, berries and fruits, roots
and tubers, insects, and carrion. When
they are available, bears will catch fishes,
and they occasionally hunt mammalian

prey, particularly newborn deer in spring.

They have the well-developed canines
of a typical carnivore, and extremely
powerful jaws with which to kill prey
and crush food. Their paws and strong
forelimbs enable them to dig, and their
sheer bulk means that they are able
to defend carrion against predators
such as wolves.
Ursus arctos

cAse sTudy

plant eaters and meat eaters

A study of brown bears in North America revealed that those which feed predominantly on meat
and fish (mainly salmon), which are comparatively rich in nutrients, are significantly heavier than
those whose diet consists mainly of plant material, which is relatively poor in nutrients: female
plant-eating bears had an average weight of 209 lb (95 kg) compared to 474 lb (215 kg) for
female meat eaters. In addition, female meat-eating brown bears had larger litters, on average,
than plant eaters, and the population density of meat-eating bears tended to be higher. Factors
other than diet may also affect bear population density, howeverfor example, human
activities, such as logging and recreational activities.

a 51 2 91 4 ft (1.72.8 m)
b Upland forests and tundra regions in North America,
northern Europe, and Asia.

c Large blonde, brown, or black bear with a wide face

and prominent mid-shoulder hump.

d 413
berry time
Brown bears on the Alaskan tundra like
to eat bright red, carbohydrate-rich
bearberries (hence their name) when
they appear in fall.

shoots and leaves

Belying their fearsome reputation,
brown bears actually consume
significant amounts of plant material,
such as plant shoots in spring and
fruits in fall.

bear, 209 lb (95 kG)

bear, 474 lb (215 kG)

food sharinG
Brown bear cubs remain with their mothers until
at least the second spring of their lives, and will
learn how to hunt, forage, and fish. This mother
bear is sharing the catch with her cub.
Gone fishinG
Brown bears exhibit a range of fishing techniques,
from waiting and watching the water before pinning
down a passing fish, to simply diving in and chasing
their prey. When fishes are plentiful, a bear will
choose the younger, more nutrient-rich fishes,
sometimes only eating the most energy-rich parts,
and discarding the remainder.

201 omnivores

occasional carnivore
Brown bear

hunting And feeding 202


Detecting prey

All organisms need energy to survive, and in order to obtain energy, most
animals have to feed on living organisms, such as plants or other animals.
Predators are animals that have evolved to hunt, capture, and feed on
other living creatures. As their prey develops ways to avoid being caught,
predators have to develop skills and behaviors that allow them to
successfully find, catch, and kill their chosen food.

Feeding strategies
Many predators specialize on one species of prey or a
few closely related speciesthe numbers of predators
and prey being an important factor in regulating both
populations. When prey is abundant, the population
of predators tends to increase. This may continue
until there is a shortage of prey, causing the numbers
of predators to fall. Large fluctuations in predator and
prey populations can occur due to factors such as
disease and weather conditions. Some predators may
take a number of species within the same size range,
while others attack anything suitable they encounter
in a particular type of habitat.




Within an animal group
there may be a range of
feeding strategies
amphibians are mostly
predatory, while most
mammal species are
herbivores. While
many species are
entirely carnivorous
or herbivorous, there
are some that are
omnivorous (use both
feeding strategies).

Cannibalism, where an animal eats members
of its own species, is widespread. Many
species practice cannibalism because of
factors such as overcrowding or food
shortage. Female tiger sharks have two
uteruses and in each, the largest of the
developing embryos will eat any smaller
embryos. Sexual cannibalism, observed in
praying mantids and some spiders, is a
special case where males are eaten during
copulation. By eating young cubs, male
lions taking over a pride bring the females
into heat so that they can mate and produce
their own litter. Sometimes cannibalism
occurs when individuals are injured or die,
as with the scavenging tadpoles shown here.

Sharks have an incredible

sense of smell and can detect
certain molecules in the water
at concentrations of only
1 part per 25 million.
predatory senses
Eye structure differs between animal
groups. Most arthropods have compound
eyes made up of multiple light-gathering
units, while vertebrates and cephalopods
have camera-like eyes that use a single
lense to focus light and form an image.

high-speed predators

To avoid being eaten, an animal
peregrine falcon
has a limited number of options.
It can avoid detection by being
camouflaged and still; it can protect
itself mechanically with spines
black mamba 20kph
or armor, or chemically with a
shortfin mako shark
noxious odor or taste; or it can try
to outrun a predator. In this case,
the need for speed has generated
an arms race over time, where hunting animals and
their quarry evolve increasingly better morphological,
physiological, and behavioral traits. Often predator and
prey become very evenly matched and success
or failure is often down to luck, surprise, or health.

Predators have evolved a range of senses appropriate

to their prey, the environment in which they live, and
their hunting strategy. In aquatic habitats, some
predatory species may simply wait for the water
currents to bring food to them. Active predators need
to look for food and can increase their chances of
success by concentrating on areas where their prey is
likely to congregate. For diurnal (day) predators, vision
is likely to be the most important sense, and their eyes
need to be forward facing to allow distance to be
judged accurately. Nocturnal predators, such as owls,
may have good hearing and eyesight. Subterranean
predators may rely exclusively on touchcave-dwelling
whip-spiders feel for their prey in total darkness using
long, slender front legs. Chemosensory organs that
detect airborne odors or waterborne chemicals enable
some predators to locate their prey at a distance.




Hearing is the ability to detect sound
vibrations transmitted through air, water,
or a solid medium. Hearing organs vary in
complexity, from simple structures to the
highly complex ears of mammals. A barn
owl can locate prey through hearing alone.






The top speed attainable by predators depends

to a large degree on the medium in which they
operate. Stooping peregrines, the fastest of all
predators, minimize wind resistance by being
streamlined and are helped in acceleration by
gravity. The high density of water restricts the
speeds that can be achieved by aquatic predators.
The cheetah is well
adapted to hunting. Its
lightweight skeleton,
large muscle mass, and
nonretractable claws give
it good acceleration and a
maximum speed of up to
70 mph (113 kph) over
short distances.

Smell is a chemoreceptive sense for
detecting odors in air. Most mammals,
especially carnivores, such as foxes,
have a well-developed sense of smell
for detecting prey. Animals give off
odors for communication purposes.
Mechanoreception is found in a wide
range of animals. It is highly developed
in spiders (such as this tarantula),
which use sensitive vibration receptors
on their legs to detect the presence
and movements of prey.
temperature sensing
The ability to visualize infrared radiation
has evolved in snakes such as boas,
pit vipers (shown here), and pythons.
Temperature-sensing organs on the
head allow these species to find warmblooded prey in complete darkness.
Echolocation (biological sonar)
has evolved in most bats,
dolphins, porpoises, and
other toothed whales. Echoes
received from high-frequency calls
are used to locate and identify prey.

silk traps
Spiders have developed the
ability to produce silk and
use it to trap insects. In
many species, the snare
takes the form of a
characteristic web, spun
across a gap in vegetation.

Many predators, whether they are active chasers

or ambushers that lie in wait, rely heavily on stealth
to be successful. Hunting of any kind expends huge
amounts of energy, and failure may not only result in
hunger but also seriously reduce the survival chances
of the predators young. Since it is likely that its prey
has also evolved good vision, often with eyes located on the sides of the
head to give a wide, nonoverlapping field of view, a predator needs to be
able to approach its target without being detected itself. Prey species may
feed in groups and large numbers of eyes mean that some can always
keep watch, making life difficult for a predator. One adaptation that helps
both predators and prey conceal themselves is camouflage. Coloring can
allow an animal to blend in with the background and disruptive markings
can help break up the body outline, which makes prey difficult to see but
also helps predators get close enough to make a kill. To increase their
chances, predators often make use of vegetation cover and move as
noiselessly as possible, keeping low to the ground as they stalk.
stalking jaguar
Hunting in the dense rain forest of Central and
South America, the jaguar takes a wide range of
prey, including caimans, anaconda, peccaries, and
capybaras. It has a very powerful bite, allowing it
to crush skulls and shatter turtle shells.

Many predators are opportunists. Rather
than targeting any particular species or
type of prey, they simply move around their
preferred habitat looking for suitable food.
Hedgehogs forage on the ground and
among leaf litter, where they find
earthworms, insects and their larvae,
slugs, and even small vertebrates.

203 predation


Several snakes have
evolved lures to entice
prey closer. Often the
lure is wiggled around,
its color contrasting
strongly with the rest
of the predators body.

A pack of wolves attack a group of musk
oxen, who will attempt to form a defensive
circle with their horns facing outward.

Cooperative hunting
In some species, cooperation enables members of a
group to obtain more food than they would be able to
by hunting alone. Cooperative hunting is more likely
to arise if individuals are related to each other and is
an important element in the evolution of social groups.
Cooperative hunting has many benets: it increases
foraging success, affords better protection from rivals
or enemies, allows larger, better-defended prey to be


targeted, and reduces the risk of injury to individuals.

In African hunting dogs, the size of the pack determines
the size of the prey. Small packs will select impalas and
small antelopes, while larger packs are able to kill species
such as wildebeests. Cooperative hunting has been
studied in vertebrates such as chimpanzees, dogs, lions,
hyenas, orcas, porpoises, sharks, shes, and birds.
Among invertebrates, social insects such as army and
driver ants are well known for this feeding strategy.


Predators have evolved an array of weapons to seize and

kill prey. The most common weapons among vertebrates
are teeth and claws. Mammalian carnivores have
enlarged canine teeth for killing, and their carnassial
teeth mesh together to tear through esh. The claws of
carnivores are large and curved for catching and holding
prey. Fish-eating species typically have a large number
of sharp, pointed teeth for securing slippery prey. Birds
that dive for shes have sharp, pointed bills with
backward-facing serrations for gripping. Birds of prey
have large talons for holding prey and strong, hooked
bills for tearing it apart. Invertebrates have a great
variety of weapons, including venomous stingers.

groove (or hollow

fang) down which
venom flows
teeth curve



lower jaw






The hyena has a massive skull with relatively short
jaws, giving it a powerful grip. In most mammalian
carnivores, the carnassial teeth are sharp, shearing
flesh before it is swallowed. In contrast, a snakes
skull is delicate, and the loosely articulated, flexible
jaws, open very wide. In vipers, killing is done using
toxic venom injected into the prey by two fangs.

Three forward-pointing
talons and one
talon ensure that this
red-backed hawk gets
a firm grip on its prey.

Scorpions combine claw
strength and venom to
subdue and kill prey.
Although this fat-tailed
scorpion has relatively
slender claws, its potent
venom allows it to easily
overcome a small reptile.

Prey may be eaten in a number of ways depending

on the predator. With very large predators consuming
large numbers of small prey, such as baleen whales
feeding on krill, there is no need for preprocessing and
the food is simply swallowed. In contrast, snakes are
adapted to eating prey as large as themselves. Since
they cannot chew it, it is swallowed whole and digested.
Most predators cut their food into pieces in order to eat
the most nutritious parts. In other animals, such as
starsh, digestion takes place outside the body.
The mouthparts of many
flies are adapted for
sponging and licking.
The house fly cannot
eat solid food so has to
liquefy it before it can be
consumed. To do this it
regurgitates part of its
previous meal, as well
as salivary enzymes,
before feeding.

Killing techniques
Predators kill in various ways depending on the size and nature of
each species. Dismemberment, strangulation, drowning, impaling, and
poisoning are the main methods. Large carnivores, such as tigers, kill their
prey as quickly as possible, avoiding personal injury or prey escape. Small
prey is bitten on the back of the neck, breaking the spinal cord and
severing the blood vessels. Death is often
instantaneous. Larger animals are
Some species are not just eaten by one
seized by the throat and held until
predator but by a whole range of predators,
they die of asphyxiation. Toxic
each using a different technique. Although
they have tough, hinged shells and strong
salivary secretions have evolved
muscles to keep them shut, bivalve
in many animals, including
mollusks are a favorite food of many
snakes, spiders, jellysh,
animals, including small sharks,
walruses, squid, and many shorebirds.
and even some mollusks.


Like all snakes, the
eyelash viper cannot
chew its food and
swallows it whole.
Once caught on the
back-curved teeth, the
prey is propelled slowly
down the throat in a
ratchetlike fashion.


Young predators such as
cats that play with their
prey before killing it are
thought to be practicing
their hunting and preyhandling techniques.
This polar bear is tossing
a piece of meat into the
air before consumption.

The shore crab uses its strong claws

to crack and pry open bivalves.

A moon snail drills a hole in the

preys shell, before injecting enzymes
to dissolve the contents.

An assassin bug picks a
weak spot in its preys
exoskeleton, such as the
soft, flexible membranes
between plates of cuticle.
Its sharp mouthparts stab
through the surface,
injecting toxic, proteindissolving enzymes. Once
liquefied, the body is
sucked up by the bug.

American oystercatchers probe

into mud and either pry or
hammer shells open.


The red-backed shrike
stores food it cannot eat
immediately. It makes a
larder by impaling items
on thorns or wedging
them in forked twigs. This
behavior allows the bird
to hoard prey when it is
abundant, and to handle
larger items such as
small lizards.
This common starsh pulls the shell
open and everts its stomach inside.

Sea otters eat oating on their backs and

often use rocks to bash open the shells.


Eating prey


bell has

Lethal sting
Box jellysh
The box jellysh uses the venom in its
stinging tentacles not just for defensive
purposes but also for killing prey. It
belongs to the class Cubozoa, and
although cubozoans are not true jellysh,
they are closely related. The box jellyshs
transparent body makes it all but
invisible to prey as it oats in the
water. Capable of moving at
speeds of up to 4 knots
(7.2 kph) using a

tentacles may reach

10 ft (3 m) in length

form of jet propulsion, it is a formidable

predator. It has 24 small eyes arranged in
four groups of six on each side of its body.
Most of these eyes are simple organs,
responding only to dark and light, but one
pair of eyes in each cluster has the
capacity to form images and may guide
the animal toward its victim. Its usual
quarry, small shes and crustaceans, are
stunned and paralyzed instantaneously
when they come into contact with the long
tentacles. As many as 15 tentacles, each
armed with thousands of stinging cells,
trail from the corners of the body. The box
jellysh is among the worlds most dangerous
venomous species and poses a signicant
threat to human swimmers and divers. Its
stings can pierce esh, crustacean cuticle,
and even mollusk shell. Excruciatingly
painful, they can cause heart failure,
although shock leading to
drowning is the most
common cause of
death in human


About 5,000 specialized stinging cells,
or nematocysts, are ranged along each
tentacle. Each cell contains a lament
armed with barbs. When triggered by
contact with its prey, the laments are
released explosively. One of the fastest
cellular processes ever recorded
in nature, this turns the lament
and barbs inside out with the
same energy as that of a
small-caliber bullet.
filament discharged
shaft inverted
operculum (hinged lid)

each tentacle
has about 5,000
stinging cells



a 10in934 ft (0.253 m) including tentacles

f Ocean waters off northern Australia, Vietnam,

Papua New Guinea, and the Philippines.

c Translucent jellyfish with pale purplish blue tinge.

Camouflaged hunter
Giant cuttlesh

Glue guns
Velvet worm
This species of velvet worm emerges after
dark to hunt the forest oor for crickets,
termites, worms, and insect larvae.
It uses its antennae to feel for prey, but has
a pair of simple eyes and can aim its glue
guns at objects up to 12 in (30 cm) away.
The mucuslike glue, which is produced by
the large slime glands that run along the
length of its body, can make up a tenth of
the weight of the animal. It is squirted from
holes at the tip of a pair of modied legs
called the oral papilla and once expelled
hardens quickly. Once the prey is trapped,
the velvet worm bites it and injects it with
salivary enzymes. The velvet worm has
curved jaws which it uses to bite off
softened parts off the prey and suck
up dissolved uids. Consuming a large
prey item may take all night.
Macroperipatus acacioi

a 41 2 in (12 cm)
f Rain forest in Brazil.
c Soft-bodied and wormlike with small surface warts

and numerous pairs of stumpy, unjointed legs.

venom has quickly

killed prey before
it damages box
jellyfishs tentacles

Speared and held fast
by hundreds of stinging
cells, this shrimp was unable to
escape before the box jellyfishs
venom took effect. The tentacle will
draw the shrimp to the underside of the
body where the mouth will engulf it whole.

Chironex fleckeri

The giant cuttlesh, the worlds largest,

stalks or ambushes its prey. Cuttlesh can
change their color and skin texture to
match their background since their skin
contains up to 200 pigment cells called
chromatophores per square millimeter.
There are three types of chromatophores
yellow, red-orange, and brown-black
and below these, reecting cells called
iridophores provide shades of blue and

Giant cuttlefish
can wiggle their
tentacles to
lure prey.

green. Cuttlesh have excellent binocular

vision and can judge distance well. Fishes
and crustaceans are snared by two long
tentacles covered in suction pads. Once
brought to the mouth, the prey is bitten
and toxic saliva injected. It is then ripped
apart by a sharp, parrotlike beak.
When prey is spotted, a giant cuttlefish turns
toward it and, holding its arms together
to guide it, launches its tentacles
with deadly speed
and accuracy.


Sepia apama

a Up to 5 ft (1.5 m)
f Rocky ledges on reefs and among sea grass off the

southern coast of Australia, including Tasmania.

c Soft elongate body with an internal bone and skirtlike

fin. Eight arms and two much longer tentacles.

d 342


Pinned down
Horned helmet shell
Also called the giant horned helmet shell
or conch, this marine snail hunts and
eats sea urchins. Males are smaller than
females and have fewer, but larger and
blunter knobs or horns. The front of the
shell has an upturned groove called the
siphonal notch along which a siphon lies
and projects into the water. It is through
this siphon that the snail tastes the
water for the chemical scent trails left by
wandering sea urchins and other types of
echinoderms, their principal food. During
the day, the horned helmet shell lies buried
in shallow water with just the tips of its
horns exposed, but when darkness falls

Poison dart
Cone shells
These sea snails hunt marine worms, other
mollusks, and even shes, paralyzing them
with the venom from a poison gland that is
injected by a harpoon-shaped dart. The
venom is very fast acting, causing almost
instant paralysis. Alerted by the smell of

it moves out of its hiding place to hunt for

food. Once it has closed in on its prey, it
raised its body and pins down its victim
using its large muscular foot. Despite its
size and weight, it is unharmed by the
urchins sharp spines.

Cassis cornuta

a Up to 15 2 in (40 cm)
f Indian Ocean from East Africa to

Australia and Pacific Ocean east to Hawaii.

c Knobbed shell with peach-colored

aperture. Males are larger than females.


>>01 The snail locates its prey by detecting its scent trail.
>>02 It raises its body by flexing its foot. >>03 The snail
moves over the urchin before pinning it down. >>04 It then
secretes mucus to dissolve a hole in the urchins shell.
Using a toothed radula, or tongue, it enlarges the hole
through which it can extract the urchins internal organs.

prey nearby, the snail extends its

proboscis, and when contact is
made it res a harpoon with
explosive speed into the body
of the prey. The victim remains
attached so it can be drawn back
into the proboscis, which expands
to engulf it. Indigestible parts of
the prey and the used harpoon
are later expelled.
The snails proboscis emerges from
the opening at the end of its shell.
Just behind its tip lies a hollow, barbed
harpoon that it uses to impale its prey.
Family Conidae

a Up to 10 in (25 cm)
f Warm tropical seas worldwide.
c Brightly colored, roughly conical shell

with an aperture running its full length.

Ordgarius magnificus
spider hangs
from foliage

a Legspan up to 1 in (2.5 cm)

f In eucalyptus trees and shrubby vegetation on the east

coast of Queensland and New South Wales, Australia.

c Pale spider with very broad yellow or pink-spotted

abdomen, and two distinctive dorsal bumps.

Deadly threads
Magnicent Bolas spider

Casting a net
Rufous net-casting spider

This bolas spider is a specialist nocturnal

predator of moths, sheltering in leaves tied
together with silk during the day. Like all
bolas spiders, it entices moths to their
doom by producing an imitation of their
sexual pheromone. It is thought that
this chemical is contained in the
sticky blobs of glue at the end of
the catching thread. The spider
aims and throws its sticky lure
toward a passing moth and,
if successful, pulls up the
thread, bites the prey,
and may eat it
immediately or wrap
it in silk for later

This spider hides during the day and

comes out after dark to catch nocturnal
prey such as ants, crickets, beetles, and
other spiders. It spins a small sheet of
pale blue, elastic, fuzzy silk held in a silk
framework and then holds the net open
with its rst two pairs of legs. Any
passing insect is enveloped in the net.

Flower pouncer
White crab spider

The rufous net-casting spider is also known as the
ogre-faced spider for the fact that two of its eyes
are very large and forward facing. It has excellent
vision and relies entirely on sight to detect prey.

Deinopis subrufa

a Legspan up to 5 in (13 cm)

f Wooded and garden habitats in southeast Australia

and Tasmania.

c Long, slender abdomen is light gray or pinkish brown

to dark brown. Slender sticklike legs.

These spiders hide in silken retreats spun

between leaves, or sit in or beside owers,
camouaged by a resemblance to ower
parts or buds. Like most crab spiders, this
is an ambush predator. At night it can
catch moths, but during the day catches
ies and even honey bees that visit whitish
owers such as daisies and jasmine. The
spider seizes the insect, bites it behind the
head, and holds it until it stops struggling.
Thomisus spectabilis

sticky globule is
swung toward prey

a Legspan up to 1 in (3.5 cm)

f Among vegetation and flowers in Australia.
c Whitish abdomen and some dark markings. Grayish

cephalothorax with pale banding on legs.

With its fangs sunk deep into its victims flesh, a
fishing spider begins to feed. As an adult, the leg
span of this species can reach 41 2 in (12 cm). It is
widespread in northern South America, where it
sits by pools in the rain forest, waiting for passing
prey, which it seizes with its front legs before
injecting a fast-acting toxin.


Leaping for prey

Jumping spiders

Silk pit
Funnel-web weavers
These spiders owe their common name
to the shape of their webs. They typically
make a at, sheetlike structure with a
funnel-shaped retreat in the middle or
at one side of the web. They spend the
day deep inside their protective retreat,
emerging after dark to hunt. Sitting in wait
at the entrance of their funnel, the spider
rushes out to grab any suitable prey that
walks across or ies onto the at part of
the web. After biting it and injecting it
with a fast-acting venom, the funnel-web
weaver takes the prey back to the safety
of the retreat to eat it. Some species,
called house spiders, make their webs
inside houses and outbuildings, usually
in dark corners and basements.

Jumping spiders do not make a web to

catch their food but stalk prey during the
day and move within range before leaping
to seize it. They have excellent binocular
vision with the middle pair of front-facing
eyes greatly enlarged. Not only is their
eyesight extremely acute, but, unlike other
spiders, they can move the back of their
main eyes inside their head so that they
can look from side to side without moving.
When hunting, these spiders attach a
safety thread to a xed point.

Family Salticidae

Jumping spiders can jump 5060
times their own body length. Their
legs extend not by muscular
action but by having fluid
forced rapidly into

a Legspan up to 5 8 in (1.8 cm)

f A wide range of habitats worldwide.
c Generally compact-bodied spider with flat-fronted

head and four forward-facing eyes. Often colorful and

brightly marked.

Decorated trap
Decoy spider
This decoy spider decorates its vertical
orb web with conspicuous patterns of silk,
called stabilimenta, which incorporate the
remains of its prey, its own shed skins, and
even plant and fungal material. Studies
have shown that by decorating their webs
decoy spiders may attract or intercept
more prey, although the decorations
may have other functions. It has been
suggested that these obvious patterns

Family Agelenidae

a Legspan up to 3 in (8 cm)
f Among a wide range of vegetation or inside

buildings worldwide.

c Hairy bodies and long, hairy legs. Drab in color, but

abdomens may have dark bars, chevrons, or spots.

>>01 Using its acute vision and a specialized
location technique, the mantis calculates the
exact distance, speed, and direction required
to snatch its prey. >>02 The front legs are
fully extended before the tibiae are flexed
around the prey in a vicelike grip. >>03 The
mantis retracts its legs and brings its
prey up to its mouth. The strike has lasted
less than 100 milliseconds.




may prevent birds and large insects from

ying through the webs by accident, or
that sitting at the center of its web, legs
folded, the decoy spider is camouaged
from predators. It is also possible that the
web debris advertises the success of
a female spider to potential mates.
Cyclosa insulana

a Legspan up to in (2 cm)
f Forests and yards from the Mediterranean

to Southeast Asia and northern Australia.

c Drably-colored spider with a misshapen,

knob-ended abdomen.

Extending mouth
Lesser emperor dragony

These large centipedes are prone to

drying out and typically forage away from
sunlight, after dark. Their prey comprises
insects, such as crickets and cockroaches,
and vertebrates such as birds, lizards, and
mice. They mainly feed on the ground
among leaf litter and under stones, but
are now known to be able to
climb up the walls of caves,
where they prey on
roosting bats, which
can be up to twice
their own mass. Despite their considerable
size, giant centipedes are extremely fastmoving and agile and can move with
ease over rocks vertically and even upside
down. The rst pair of legs has been
modied into stout, sharp claws that inject
powerful venom from poison glands. The
venom is usually injected near the head of
their prey, where it works quickly, and
while the prey struggles, the centipede
wraps its other legs around it.

As with all dragonies, the nymphs of the

lesser emperor dragony live in lakes and
ponds among vegetation and bottom
debris, where they hunt for prey which
they catch using a concertina-like lower
mouthparts (see panel, right). Their gills
are located within the rectal chamber and,
if required, water can be forced out at high
speed for a jet-propelled escape from
potential enemies.

Scolopendra gigantea

a Up to 14 in (35 cm)
f Forest in Central and South America.
c Robust flattened bodies with reddish brown segments

each with a pair of yellowish legs. Head has a pair of

segmented antennae and strong mandibles.

Sudden ambush
Common praying mantis
Like all other mantids, the common praying
mantis has a highly specialized predatory
lifestyle. A skilled ambusher, it remains
motionless and relies on its cryptic green
or brown coloration to avoid being seen. It
has a distinctive triangular head that is very
mobile and has a pair of large compound
eyes, which face forward to provide it true

Anax parthenope

a 234 in (7 cm)
f Lakes and ponds in Europe and Asia.
c Greenish brown abdomen with broad blue bands at

base, and pale greenish brown thorax.

A bat retuning from a
nocturnal foraging trip
has been plucked out of
the air at the entrance
of a cave. Giant
centipedes can hang
from the ceiling or an
overhang, maintaining
a firm grip with at least
five pairs of legs.


Climbing the
Giant centipede


The aquatic nymphs (or
naiads) of dragonies are
highly predatory and have
specialized mouthparts
(mandibles) for catching prey.
The mouth is long, hinged,
and prehensile, with a pair of
hooked palps, and folds back
on itself. It can be shot
forward using muscular
action and hydraulic pressure
in 25 milliseconds or less.
>>01 A passing fish has alerted this dragonfly nymph to its presence.
The nymph watches it for a short distance. >>02 When ready, the
nymph launches itself upward and extends the lower mouthpart forward
from underneath the head. >>03 Impaled on either side
by the sharp hooks on the palps, the fish is brought to the mandibles.




binocular vision. It gauges distance to its

prey by moving its head to measure the
preys apparent movement relative to
its background. Known as binocular
triangulation, this technique is
widespread among vertebrates
but much less common
among invertebrates. There
are many modications
of the mantiss body
that make it a superb
hunter. The rst

muscular front

mantiss tibia is
folded back along
its femur to hold
prey fast before it
is devoured

segment of the thorax, which carries

the specialized front legs, is very long.
Together with the elongated upper
segment of the front legs, this gives the
mantis a very broad reach. The front femur
is greatly enlarged to house the muscles
that operate the tibia, and has rows of
sharp spines on its inner surface. The front
tibia is also spined and folds back like a
jackknife to mesh with the spines on the
femur, making a formidable trap. The
middle and hind pair of legs are used for
walking and holding onto vegetation. The
common praying mantis is mainly active
during the day and eats a wide range of
insects, spiders, and other arthropods.
When prey is caught and subdued, the
mantis uses its tough jaws to slice through
tissue and chitin with equal ease.

Mantis religiosa

a Up to 234 in (7 cm)
b In all kinds of vegetation in Europe; introduced

to North America.

c Elongate, green or brown insect with large,

spiny front legs and distinctive triangular head.

The prey is impaled and
held by the front legs,
so there is no need for
venom, and a mantis
eats its food alive.
Generally the whole
victim is eaten, leaving
only fragments.


Aquatic assassin
Giant water bug nymphs
Giant water bugs are erce predators both
as adults and as nymphs. Nymphs have
large bulging eyes and either lie in wait
for prey to pass close enough to lunge
forward and grab it with their specially
modied front legs, or hunt actively using
their middle and hind legs to swim. Once
the prey is caught, the bug uses its sharp

Surface predator
Common pondskater
These gregarious bugs are well adapted
for catching small insects trapped by the
lm at the surface of fresh water. The front
legs are short and used to grasp prey as
the pondskater eats, while the middle
and hind legs are very long, and splay out
to support the insect on the water and
propel it across the surface at great speed.
Pondskaters are covered in dense,
water-repellent hairs, and the front legs
have ripple-sensitive hairs that enable
them to detect and locate the movement
of their prey.
Gerris lacustris

a 1 2 in (1.5 cm)
b On still or slow-moving water in Europe.
c Slender body, brownish above and silvery gray

beneath, with long middle and hind legs.

mouthparts to stab it, usually in the neck,

and inject highly toxic saliva, which
paralyzes the victim. Similar in composition
to some snake venom, the toxin dissolves
the preys esh so that the liqueed meal
can then be sucked up by the bug.
Although the larvae of aquatic insects form
a large part of the diet of giant water bugs,
surprisingly, they will also readily attack
craysh, tadpoles, frogs, shes, and even
small water birds, all prey that are much
larger than themselves.

This giant water bug nymph has successfully
ambushed a frog by attacking it from below,
injecting toxic venom, and keeping clear of
the amphibians powerful hind legs.

Family Belostomatidae

a Up to 6 in (15 cm)
b Freshwater streams and ponds worldwide, especially

the Americas, southern Africa, and Southeast Asia.

c Streamlined body with strong grasping front legs.

The saliva of giant water bug

nymphs is similar to the venom of
some snakes. It is extremely toxic
and paralyzes the bugs victims.

Open jaws
Giant antlion larva

Palpares immensus

Giant antlion larvae bury themselves in

coarse sand, with just the top of their
heads and open jaws showing above the
surface, while they wait for insects and
other prey to ambush. Sensitive hairs and
relatively good eyesight tell the antlion
when to strike. Anchored deep in the sand,
it can subdue large prey. The toothed,
sickle-shaped jaws are two hollow tubes
through which salivary enzymes can be

which has very large jaws armed with sharp teeth.

a 2 in (5 cm)
b Sandy areas among long grass in southern Africa.
c Fat-bodied with a slender neck and squarish head,

injected into the prey to paralyze it and

dissolve its internal organs. The resulting
soup is then sucked back up and eaten.
Some antlion larvae build special conical
pits in loose sand. They then ick sand
grains at passing insects to knock them
down into the bottom of the pit where
they are seized and eaten.

Once the toothed jaws of the antlion larva
have snapped shut, crushing the preys
body, there is no escape. It will quickly be
dragged under the surface and eaten.

Often called glow worms,

the larvae of this gnat live in
a mucus tube supported by
a loose scaffolding of silken
threads. From this structure
hang as many as 50 vertical
threads coated in small beadlets
of a sticky, gluelike substance.
To attract ying insects such as
mosquitoes, midges, mayies,
caddisies, and even beetles
to their sticky snares, the larvae
produce a soft blue-green
glow from the rear ends of
their bodies. When an insect
becomes trapped in the glue,
the larva moves slowly along
one of the horizontal parts of the
web to the end of the thread
and begins to reel the victim in.
The gnat larvae have to live in very
sheltered locations as the slightest
gust of wind would tangle the sticky
beadlets on the threads together,
making the trap useless.

Arachnocampa tasmaniensis

a Up to 1 4 in (3 cm)
b Caves, overhangs, and deep gullies
in Tasmania.

c Soft-bodied, pale, maggotlike larva.


Sticky snare
Fungus gnat larva

sensitive antennae
large compound eye gives
accurate eyesight

Bee killer
Asian giant hornet
Asian giant hornets hunt and
kill a wide range of insect
prey, including other
hornets. Able to cover
relatively long distances,
the hornet carries its prey back to its nest,
where it uses its strong jaws to butcher the
victim. It is then fed to the developing
hornet larvae. However, the adults do
not feed on what they catch, but are
instead fed by their own larvae, which
regurgitate a rich mixture of amino
acids for them to drink. When Asian
giant hornets attack bee colonies, it is
not the adult worker bees they carry off,
but the soft-bodied bee larvae. Just a
handful of hornets can decimate a bee
colony in a short time, an attack for which
the introduced commercial honey bee has

broad wings
powerful mandibles

no defense. However it
has been discovered that
Japanese populations
of the Asian honey bee have evolved a
unique thermal-execution technique to
deal with such an attack. Hundreds of
worker bees form a tight ball around the
invaders, and the intense heat generated
by their wing muscles literally burns the
hornets to death (see p.313).


This species can fly
at more than 12 mph
(20 kph). In addition
to powerful venom, it
also has strong jaws.

Hairy trap
Allomerus ant

Vespa mandarina

a Up to 2 in (5 cm)
b Mostly upland areas in Japan, China, Korea, Taiwan,

India, and Nepal.

c Large-bodied hornet with broad orange head, dark

thorax, and yellow and black banded abdomen.

The stem of the ant plant Hirtella physophora
is covered by a secondary surface built entirely
by the hiding ants.

These ants have evolved a unique

mechanism for capturing large prey that
might otherwise escape them by jumping
or ying away. First, they trim away some
of the hairs of a host plant stem, leaving
pillars of hairs. These are used to support
a spongy platform that they make using
regurgitated material combined with the
trimmed hairs. The platform is disguised
as a hairy stem and made with many small
holes so that the ants can hide beneath.
The construction is reinforced by a species
of sooty mold that is controlled by the ants.
Once the trap has been set, the ants hide
below, with their jaws open, ready to catch
any prey that passes along the stem.

>>01 When large prey such as a grasshopper
walks on the surface of the ants trap, its legs
are grabbed from below, and it is then
gradually stretched like a victim on a medieval
torture rack. >>02 Once immobilized, the
grasshopper is stung. >>03 The prey is then
carried along to the ants home inside a leaf
pouch, where it will be cut up and eaten.

Allomerus decemarticulatus

a 116 in (2 mm)
b Rain forest in South America.
c Yellowish orange, smooth-bodied ant.

Dwarfed by the
supermajor worker,
swarms of minor
workers aid their much
larger counterpart.

Pheidologeton affinis

a Up to 1 2 in (1 cm)
b Forest and grassland in Southeast Asia
c Shiny yellowish brown to brown head, thorax

and abdomen, and yellowish legs.



Super-sized raider
Marauder ants

Path of destruction
Driver ants

In this species there is a great variation

in size between different castes. The
workers fall roughly into three sizes: very
small workers, known as minor workers;
major workers; and large ants known as
supermajor workers, which may be several
hundred times heavier than
the smallest workers of the
colony. This extreme range
of worker sizes allows these
ants to utilize a much larger
range of prey and to defend
themselves against attack.
In large numbers, marauder
ants can even attack the nests of green
turtles. Marauder ants go out on foraging
raids for prey to feed to their developing
larvae but tend to have more permanent
nest arrangements than other raiding ants.

Also known as safari ants, these ants form

colonies that can comprise many millions
of individuals. They may stay in one place
for anything from a week to several
months. When local food supplies start
to diminish, the ants move off in a large
column, consuming anything they
encounter. During this nomadic phase,
the workers carry their developing brood
in their jaws. Although insects and other
invertebrates are their main prey, larger
animals such as vertebrates can be
overwhelmed and if unable to escape,
will be hacked to pieces and eaten.
Dorylus species

a Up to 1 in (2.5 cm)
b Savanna and woodland in Africa and East Asia.
c Brown to black bodies with paler legs.


Bigger soldier ants,
which are armed with
large, curved and
toothed jaws, patrol
the outside edges of
the perpetually moving
column and are
constantly on guard
for potential attackers.

The white shark, a formidable predator

also known as the great white, typically
employs a lightning-fast strike from below
to disable large prey. The blow can seriously
incapacitate or even kill prey in an instant.
Despite its size, the white shark is
effectively camouflaged by countershading.
Its body is gray or bluish brown on top so
that prey swimming near the surface find it
hard to see the shark when looking down
into the water beneath them. Similarly, the

sharks underside is pale so that when

its prey is below, the sharks silhouette
against the sky is minimized.
When hunting, white sharks are able to
maintain their body temperature, especially
that of their brain, and use a range of
senseselectroreception, scent, hearing,
and visionto locate prey. They will often
have an exploratory bite to test if an item of
prey is suitable. Juveniles usually eat rays,
other sharks, and squid, but adults hunt
seals, sea lions, dolphins, and some large
fishes, such as tuna. Although this shark is
a top predator, occasionally it will scavenge
carrion such as whale carcasses.


25 mph The

speed reached by
white sharks when
they attack prey
from below.

edges used
for slicing

jaws up to 31 4 ft
(1 m) wide

Carcharodon carcharias

a Commonly up to 20 ft (6 m), but may be longer.

b Found mainly in coastal waters where prey is

abundant. Worldwide in temperate waters.

c Robust, torpedo-shaped body. Head has a conical

snout and small, dark eyes. Pectoral and first dorsal fins
are large, and two tail lobes are similar in size.

humAn ImpAcT

attacks on people
The recent decline in white shark populations
reflects the fact that the losers in most shark
human encounters are the sharks. As humans
spend more time in the sea, they inevitably
encroach on the habitats of sharks, so this
enforced interaction has also resulted in
increased numbers of shark attacks.
However, the prevalent perception that
sharksespecially the white sharkare
man-eaters is contradicted by the fact that
a decreasing proportion of shark attacks
has resulted in fatalities.
increasing survival rates
While the number of white shark attacks on
humans increased during the 20th century, the
proportion of fatalities dropped considerably,
probably due to improved medical help.



attack from beneath

White shark


10 20 30 40 50


rows of teeth
grow forward
to replace
lost teeth

jaws of death
The large triangular teeth of the upper jaw
have serrations on both sides and are used
for slicing through flesh. The teeth of the
lower jaw are much smaller and
help secure prey.

full breach
The sheer force of its attack has
carried this female white shark clear
of the water. Having spiraled through
the air, she re-enters the water with
her prey still gripped in her mouth.

impact from below

>>01 This shark has stalked and attacked
a seal from below, launching it out of the
water. >>02 Although the seal has avoided the
sharks grasp, its body has taken the full force of
the strike. >>03 The injured seal falls back into
the water. The shark may attack the seal
repeatedly until it is weak enough to be eaten.


Like other hammerheads, the scalloped

hammerhead sharks most distinctive
feature is its broad, at head. The most
likely explanation for this bizarre shape
concerns the capture of its prey. All sharks

Night hunting
Sand tiger sharks
Also known as the gray nurse shark or raggedtoothed shark, this large, slow-moving species feeds
mainly on shes, skates, and rays as well as large
crustaceans and cephalopods. During the day, it
typically hides in caves and reefs, sometimes
swimming in small groups very close to the bottom.
At dusk it moves into the open to hunt for food.

Sand tiger sharks are often solitary, but
when food is abundant they can feed
together. Like all sharks, they have
electroreceptive organs to locate prey.

have special sense organs, called the

ampullae of Lorenzini, capable of detecting
weak electrical elds emitted by other
animals. These organs form a system of
jelly-lled canals in the head and snout
connected to the outside via small pores. In
hammerhead sharks, the underside of the
hammer is particularly well endowed with
these organs, and the increase in area due
to the shape of the head may help them
detect bottom-living prey more effectively.

It is thought that sharks
can detect electrical
signals as low as 15billionths of a volt from
the movement of prey.
Sphyrna lewini

Carcharias taurus

a Up to 101 2 ft (3.2 m)
b Typically in shallow sandy areas

and reefs, almost worldwide.

c Stout, grayish brown body with flattened

head, conical snout and prominent teeth.

a Up to 13 34 ft (4.2 m)
b Warm temperate to tropical coastal

waters worldwide.

c Brownish gray above and pale below.

The hammer-shaped head has a wavy
or scalloped front edge.

Hidden angler
Hairy frogsh

Water cannon
Banded archersh

Also known as the striated or striped frogsh, the

hairy frogsh is a stealth predator that lives on the
seaoor hidden among rocks, weed, and coral.
Concealed by highly effective camouage, it draws
prey toward it with a lure. Its coloration is very
variable, and its body is covered with long and
branching skin tufts that completely obscure the
outline of the sh. The tubelike openings of the gills
are hidden just below the pectoral ns, and to make
their disguise even more effective they will hide
among urchins and sponges, and in coral crevices.
The hairy frogshs stalked lure is a modied dorsal spine
that can be bent backward along the head when not being
used to attract prey. The lure can even regrow if it gets damaged
or bitten off. Frogshes do not always need to use their lure, since
they are so well camouaged against the substrate that small
shes will sometimes even mistake them for a place to shelter.

Although the banded archersh catches aquatic prey such as

shrimps and worms, it is best known for the unique way it catches
prey out of water. The sh uses its mouth to squirt a narrow,
powerful jet of water to knock insects from the vegetation above
it. The head is narrow with the eyes close to the snout, giving it
binocular vision. When the sh spots potential prey, it sticks its
snout out of the water and presses its tongue onto a groove that
runs along the roof of its mouth. By rapidly closing its gill covers,
a stream of water is forced along the groove and out through the
small aperture at the end. Adults are very accurate and can shoot
a number of times in quick
succession to dislodge prey,
Toxotes jaculatrix
which is then swallowed,
a Up to 151 2 in (40 cm)
sometimes before it reaches
b Mainly in mangrove swamps of
the water. If prey is close to
Southeast Asia, India, Australia, and the
the water, the archersh
may jump out of the water
c Silver-bodied and laterally compressed
with dark vertical bands.
to catch it.

The speed of the frogfishs gape-and-snatch
movement has been measured at an extraordinary
1/6,000th second, making it among the fastest
known predatory movements of any vertebrate.
Antennarius striatus

a Up to 10 in (25 cm)
b All subtropical oceans, to depths of 33656 ft

(10200 m).

c Variable coloration, often light yellow, orange, green,

gray, or brown with black stripes or elongate blotches.

Eyes have prominent radiating lines.

location of

of refraction

enlarged dorsal
spine has appearance
of weed


bait at end
of stalk
swims just
below surface


Frogfishes sit absolutely
motionless on the seabed,
camouflaged against the
substrate. This keeps
them hidden from
predators but also
accentuates the movement
of the lure, which they
twitch to attract prey.

An archerfishs aim
improves with age.
Adults can spit a jet of
water more than 6 ft
(2 m). They can also
judge the size of the
prey and therefore how
much water is needed
to knock it down.

As light passes from
air to water it bends
or refracts. The
archerfish is able
to compensate for
this refraction and
shoot at the true
location of its prey.


Electrical sensing
Scalloped hammerhead

Also known as the copper shark due to its color
in sunlight, the bronze whaler shark can grow
to over 9 ft (3 m) in length. It feeds on other
sharks, squid, and bottom-living fishes but is often
attracted by large schools of fishes, such as
sardines on an annual run that takes place off the
east coast of South Africa between May and July.


Concealed weapon
Mosaic moray eel
Hiding in crevices and under rocky ledges,
these thick-skinned, mucus-covered eels
typically lie in wait for something to pass
close enough to be seized. They mainly
feed at night on shes, crustaceans, and
mollusks and move with a snakelike,
sinuous motion powered by their long
dorsal and anal ns. They sometimes feed
during the day and have been observed
to stalk their prey over short distances.
The eels head has a characteristic long
snout with curved jaws, armed with many
razor-sharp, needlelike teeth. A second set

of jaws (see panel, below) ensures that the

eel can secure its prey even if it only gets a
partial grip on it. Other bony shes typically
use suction to swallow prey; sudden
expansion of the mouth cavity causes
water to rush in, taking the prey along with
it, but in moray eels, the head is relatively
narrow so it would be difcult to ingest
anything but small prey using this feeding
Enchelycore ramosa

a Up to 5 ft (1.5 m)
f On rocky reefs off southeast Australia, northern

New Zealand, and the South Pacific.

c Snakelike, scaleless, yellowish gray body with a

mosaic-like pattern.


Moray eels have a second set of
jaws halfway down their throat
called the pharyngeal jaws. Prey
items are caught and held fast by
the sharp teeth on the primary
jaws, but because it is difcult to
pull food in through the narrow
mouth using suction (see above),
gill arch free of spine
another mechanism is needed.
Contraction of special muscles
brings the pharyngeal jaws into
play in a mechanism unique to
moray eels. These secondary jaws,
also bearing large, back-curved
teeth, move forward rapidly from
well behind the skull to take up a
jaw muscle
position inside the mouth. Once
anchored to
the prey is secured, other muscles
contract, pulling the pharyngeal
lower pharyngeal
jaws backward, and dragging the
jaw muscle
prey down into the esophagus.
anchored to gill arch

Although their eyes are
prominent, moray eels have
relatively poor eyesight, and
rely instead on a good sense
of smell and sensory pores on
the head to detect their prey.

Secret flashlight
Stoplight loosejaw

The stoplight loosejaws
striking appearance
remains hidden from
the crustaceans and
fishes that it feeds on.

red light is invisible

to other fishes and
helps to detect prey

Malacosteus niger

a Up to 10 in (25 cm)
f Deep areas of the Atlantic, Indian, and

Pacific Oceans.

c Scaleless, with a large head and tapering black body.


blue sensitivity of
most deep-sea
animals eyes

red sensitivity
of stoplight
loosejaws eyes

red emission of
Most deep-sea species can see
stoplight loosejaws blue-green bioluminescence (light
light organ
generated by living organisms) but

are not very sensitive to red light.

The stoplight loosejaws flashing
red photophore allows it to see its
prey close up while remaining
invisible. This species is unusual
among vertebrates in being able to
produce long-wavelength (far red)








(lower main jaw)
upper pharyngeal jaw
lower pharyngeal jaw

upper pharyngeal
jaw brought forward
lower pharyngeal jaw
brought forward

These nocturnal fish hide during
the day in dark crevices and under
submerged wood. They have
poor eyesight but have a special
electrolocating system to move
around in the darkness and find food.

This shs common name comes from the

presence of red and green photophores
(light-producing organs) on the sides of its
head, which it uses to catch prey. The
lower jaw is very long and armed with
slender fangs that curve backward.
To seize prey, the jaw can be pushed
forward in front of the head, and once prey
is snagged on the teeth, the jaw is
retracted. There is no skin between the
bones of the lower jaw, and the species
can swallow prey almost as big as itself.
light used to
attract prey


Gnathonemus petersii

a Up to 14 in (35 cm)
f In muddy, slow-moving rivers in West

and Central Africa.

c Dark fish with rectangular, flattened

body ending in a slender, forked tail.

Electric organs
Elephantnose sh
These shes live in muddy and murky
water and need a system other than
eyesight to locate and determine the
nature of nearby objects. They use a kind
of radar by producing electrical discharges
from a special organ made up of modied
muscle cells near the tail. The discharges
then generate a weak electrical eld
around the mucus-covered body of the
sh. Obstacles, prey, or other shes that
come into range of the electric eld alter its
shape according to their own conductivity,
and these changes are picked up by
electroreceptors all over the body. These
receptor cells are particularly abundant on
the head and the strange-looking,

rear electrical receptor


front electrical receptor

or schnauzenorgan

The scientific name meaning thread jaw refers
to the fingerlike sensory organ on the bottom jaw,
which is used to feel for worms, insects, and
crustaceans in the bottom sediment.
downward-curved snout or schnauzenorgan,
which the sh also uses to probe the
bottom. The brain of these shes is
enlarged to process the complex
information coming from the electrolocating system, the ratio of brain to body
mass being close to that of primates.

Plankton pickers
Garden eels

Wild brown trout return from the sea to

spawn in fresh water and while there eat
a wide range of insects, worms, mollusks,
crustaceans, and even smaller shes and
frogs. They feed from the bed of the
stream, and when certain insect species
are hatching and rising to the surface, the
trout gorge themselves. They often rise
out of the water to reach ying insects.
Because of their varied feeding habits,
anglers use lifelike models of a wide range
of insects for y shing, where they try to
mimic the appearance and behavior of the
trouts natural food. Trout can be very
selective in their feeding, and shermen
have to try to imitate what the trout are
currently eating.

Garden eels have good eyesight and rely

mainly on this sense to catch zooplankton,
other invertebrates, and even small shes
that drift past in the current. They live in
colonies and typically remain in their
burrows in the sand. Even when feeding,
they only emerge partially, and always

leave one-third of their body length buried

below the surface. The burrow is coated
on the inside by mucus secreted by a
special gland in the eels tail. This
substance binds the sand grains together
and prevents the burrow from collapsing.
Also known as the Indian spaghetti eel, the
garden eel is extremely wary of predators,
and will retreat in an instant, sealing the
entrance of the burrow with a plug of
mucus until the danger has passed.

Gorgasia maculata

a Up to 28 in (70 cm)
f Sandy shallows of the western Pacific Ocean, from

the Maldives to the Solomon Islands and the Philippines.

c Pale and slender-bodied, 3 8 in (1 cm) wide.

Garden eels live gregariously in colonies known
as eel gardens (hence their name), where they
sway around like stalks of sea grass.

Salmo trutta

a Up to 41 2 ft (1.4 m)
f Rivers and streams of Europe and Asia, and the

northeast Atlantic, but widely introduced elsewhere.

c Stocky body with olive green to silvery dorsal surface

covered with darkish spots.
Using its powerful tail to jump clear of the water a
brown trout will sometimes try to catch fast-flying,
agile prey such as damselflies.

Side swipe
Atlantic sailsh
Sailsh feed on very fast-moving shes
such as mackerel and tuna, striking down
or sideways at prey from behind using their
long bill. Groups of sailsh can use their
sails to herd schools of sh. One of them
then scythes through the school, thrashing
its bill from side to side to cripple and kill
the shes, which are eaten head rst.


The characteristic sail is the large first dorsal fin,
which can run almost the entire length of the body.
The sail is usually kept down and to the side but
erected when the fish is threatened or excited.
Istiophorus albicans

a Up to 934 ft (3 m).
f Caribbean Sea, extending across Atlantic Ocean

to West Africa.

c Smoothly tapering blue body, with blackish blue fins.

Elongated upper jaw forms a pointed bill.

Unfolding tongue
Perezs frog
These frogs never venture very far from permanent bodies of
water and typically sit at the waters edge on muddy banks or
rocks to sunbathe and feed. The males have a pair of vocal sacs
and call to attract mates. They can be active by day and night,
and their diet consists primarily of insects, spiders, and other
small invertebrates, which they catch using their sticky extendable
tongues. When not in use, the tongue is folded backward toward
the throat. A row of very small teeth in the upper jaw,
as well as some small teeth in the roof of the mouth, hold prey
in place before it is swallowed. The frogs in turn are eaten by
owls and several species of aquatic birds. They are always ready
to dive into the water and hide at the slightest sign of danger.



Pelophylax perezi

a Up to 4 in (10 cm).
f Rivers and ponds in France and the
Iberian Peninsula.

c Greenish, brown, or gray with mottled

pattern with long pale dorsal stripe.

The tongue of the Perezs frog has a
sticky upper surface, and is attached
toward the front of the mouth rather
than at the rear. This, together with
their ability to jump, gives the frogs a
longer reach for capturing prey.

Frogs large, bulging

normal position
of eyes
eyes have an extra job
besides watching prey.
frog has prey
When a frog swallows
in mouth
its prey, the eye
muscles contract,
head is
pulling the eyes shut
and moving them
down through small
initial eye
openings in the skull.
The eyes take up a
space near the back of
eyes are fully
the mouth and this
action helps push the
prey is pushed
food down.
toward stomach


Fly catcher
Brown trout


The komodo dragons powerful legs
and strong claws hold down prey as
they tear off large chunks of flesh.
To accelerate its digestive
process a lizard will then
bask in the sun.

Lethal bite
Komodo dragon
Although Komodo dragons also eat carrion,
they will ambush other reptiles and small
mammals, as well as goats and deer,
usually administering a lethal bite (see
panel, right). Swallowing can be a slow
process, and so that the lizards do not
asphyxiate while eating, a breathing tube
under their tongue connects to the lungs.
A loosely hinged lower jaw and an elastic
stomach deal with meals that may be up
to 75 percent of their own bodyweight.


The mouth and teeth of Komodo dragons
are home to more than 50 species of
pathogenic bacteria. Animals that are bitten
and manage to escape will usually die from
the effects of blood poisoning. This may
work to the lizards advantage since it is
able to detect the smell of a dead animal up
to 6 miles (10 km) away. The bite of other
monitor lizards is also slightly venomous,
causing swelling and pain for many hours.

Varanus komodoensis

a 69 ft (23 m)
b Grassland and lowland forest on the Indonesian

islands of Komondo and Flores and others in between.

c Heavy bodied, grayish brown with darker markings,

very long tail and long yellow forked tongue.

d 373

River ambush
Nile crocodile
Young Nile crocodiles eat insects and
other invertebrates, and as they grow,
will take larger prey such as reptiles,
amphibians, and birds. Their ability to rest
motionless in the water with just the eyes
and the tip of the nostrils showing allows
them to get very close to unwary prey
before they launch an attack. On land, they
are rather slow moving, but can swing their
legs underneath the body for a faster gait.
In the water, they are incredibly
With a huge gape and formidable
bite, a Nile crocodile can hold onto
large prey such as zebras,
buffaloes, or wildebeest and drags
them into the water to drown.

swift, using their powerful body and tail to

propel themselves forward at bursts of up
to 18 mph (30 kph). Adults usually eat
sh, but will lie in wait for animals to
come down to the waters edge to drink.
Sometimes they gather in numbers at river
crossings where groups of migrating
species such as
wildebeest have
to run a deadly
gauntlet. Since
they cannot
use their
teeth to cut

meat, crocodiles spin themselves around

in the water to tear meat from the carcass
although this is known as the death roll,
the prey is usually dead already. Nile
crocodiles can go for long periods without
eating but when food is available they can
consume up to half their own body weight
at a single meal.
Crocodylus niloticus

a Up to 20 ft (6 m)
b Waterways throughout Africa and western Madagascar.
c Large and squat, short splayed legs and long tail. Back
dark with tough scales, underside yellowish and softer.

d 401, 476

A group of hungry Nile
crocodiles converge on a
zebra victim before
ripping it apart.

19 ft The length
of some Nile
crocodiles, the
largest freshwater

This turtle lies in the water with its hooked

jaws held wide open. The inside of its
mouth, like the rest of the head, is drably
colored and patterned to blend in with its
background except for the tip of its tongue,
which is long and reddish. The turtle
wiggles its tongue around, imitating the
movement of a worm to lure shes close
enough that it can snap its jaws shut on
top of them. In addition to eating shes,
snakes, amphibians, and even other
turtles, this species will also readily eat
red tongue
resembles carrion if given the opportunity.


Tongue lure
Alligator snapping turtle

Vivid green coloring
with blue, yellow, and
white flecks give superb
camouflage, while
enlarged front teeth
enable the snake to
grasp its prey securely.
large eye with
vertical pupil

temperaturesensing pit

a worm

Even the outline of the turtles eyes is
broken up by radiating lines, so the only
conspicuous thing is its red tongue.
Macroclemys temminckii

spiked plates.

Most toxic
Inland taipan
Small mammals such as mice and rats
form this snakes prey, and it will also steal
their burrows. It gives several bites in quick
succession, and the venom is so fast acting

that the prey is limp and paralyzed almost

the instant it is bitten. The inland taipan
is the most toxic land snake in the world.
The venom is many hundreds of times more
toxic than that of a diamondback rattlesnake
and 50 times more toxic than that of the
Indian cobra. A bite can prove fatal to a
human unless antivenin is available.

Having a dark head allows the snake to warm
up rapidly up in the morning sunshine by
simply poking its head out of its burrow.

Sensing blood
Green tree python
This snake is a nonvenomous constrictor
that lives mainly in the tree canopy and
eats small mammals, such as rodents and
bats, as well as reptiles and birds. It rests
curled in loops over horizontal branches
looking out from the middle of the draped
coils. Prey is usually caught by the snake
holding onto a branch with the rear part of
its body while it strikes with its head. The
snake has special temperature-sensing
pits along the margins of the lower jaw that

enable it to detect the presence of warmblooded prey. The pits detect radiant heat
and allow the snake to track and catch
prey even in total darkness. Once prey is
caught, the snake wraps several coils of
its body around it and tightens its grip
every time its victim exhales, suffocating
it in the process.


a Up to 26 in (65 cm)
b Lakes and rivers in southern North America.
c Brown turtle with hooked jaws and shell with

Morelia viridis
Oxyuranus microlepidotus

a Up to 61 2 ft (2 m)
b Arid scrub and grassland in central Australia.
c A glossy, grayish brown snake with irregular

dark markings.

A northern cat-eyed
snake, hunting in
forest foliage, gorges
itself on frogspawn,
its preferred food. It
frequents any lying
water where frogs and
lizards lay their eggs.

a Up to 61 2 ft (2 m)
b Rain forests in New Guinea, Queensland Australia,

and some islands in Indonesia.

c Bright green snake with a broken pale stripe along

the back.

Spawn hunting
Northern cat-eyed snake

Death grip
Central American boa

Known as a cat-eyed snake because of its

slightly protruding golden-brown eyes with
catlike vertical pupils, this species will eat
adult tree frogs, toads, and lizards if it gets
the chance, but prefers to eat frogspawn
and young tadpoles. The viperlike head
has grooved fangs at the back of the
mouth, and the venom it injects is only
mildly toxic. Cat-eyed snakes hunt at dusk
and after dark around ponds, forest pools,
and streams where frogs are mating and
laying eggs. Although they do hunt on the
ground, in forested areas they will climb
among the branches of trees to look for
tree frogs. Attracted by the frogs nocturnal
mating calls, the snake silently slithers
among the foliage to feast on masses of
their spawn. These snakes are not very
abundant; however, to compensate for
this, mated female cat-eyed snakes can
store sperm for several years.

The general background color and

patterning of these solitary snakes serves
as good camouage against the forest
oor and among trees. Boa constrictors
eat rodents, bats, and other mammals as
well as birds and lizards, and when fully
grown can tackle capybaras,
monkeys, and even
wild pigs. To help
them locate prey,
there are some
simple heatsensitive scales on
the head, and they
have a good sense of
smell. When catching
bats, they hang from
trees or cave entrances
where the bats roost and
snatch them out of the air
as they y past. The mouth
has curved teeth to hold the
prey as the body coils
around it. Once the prey has
been killed and swallowed
whole, it may take several
weeks for the snake to digest
it completely.

Leptodeira septentrionalis

a Up to 35 in (90 cm)
b Lives in forested areas in the southern US, Mexico,
and Central America.

c Triangular head, dark bands on paler grayish brown,

narrow body.



Seen through a thermal imaging camera, a rodent
stands out clearly from the cool background. Prey
may seem just as obvious to the green tree python.

Boa constrictor imperator

a Up to 143 4 ft (4.5 m)
b Forest, savanna, cultivated areas, and

mangrove swamps in Central and South America.

c Stout-bodied snake with flattened head and

dark markings.

Boa constrictors are
nonvenomous and kill
by asphyxiating their prey,
squeezing them slowly to
death in their powerful coils.

The tongue of a panther
chameleon is extended by
a unique system that
catapults it toward prey at
speeds in excess of 16 ft
(5 m) per second. The
sticky end wraps around
the prey, and the tongue is
retracted into the mouth.


Flexible jaws
Common egg-eater snake
These snakes live in forests and wooded
areas populated by nesting birds. A good
sense of smell allows them to detect
whether or not an egg is rotten before they
try to eat it. A highly exible neck enables

them to take even large eggs into their

mouths whole, before the shell is broken
by teethlike projections from the spinal
bones. The snake then swallows the liquid
contents and regurgitates the fragments of
broken shell. Egg-eating snakes can fast
for long periods. When food is plentiful,
they gorge themselves and their metabolic
rate increases, as does the diameter of

the liver and small intestine. When

threatened, the common egg-eater snake
can generate a loud hissing noise by
repeatedly coiling and uncoiling its body,
which rubs its lateral scales together. They
will also strike out open-mouthed, even
though they have no teeth to attack with.
Dasypeltis scabra

This large eagle

occupies the same
ecological niche as
the bald eagle in
North America.

a Up to 3 ft 3 in (1 m)
b Forest and woodland in sub-Saharan Africa.
c Slender snake with grayish brown body and
rough scales.

>>01 By dislocating its jaw, the snake begins
to consume an egg that is much larger than its
head. >>02 Once it has taken the egg into its
throat, the snake begins to close its mouth
around it. >>03 Muscular contractions press
bony projections of the spine on to the egg to
break it. The snake then swallows the contents.

Flightless forager
Stewart Island brown kiwi

Dive bombing
Peregrine falcon

This bird is a subspecies of the Tokoeka,

a New Zealand kiwi. Like other kiwis, the
Stewart Island brown kiwi is ightless and
well adapted to terrestrial living and
foraging. It has large, powerful legs with
claws to scrape and dig, and, unusually
for a bird, foot pads that enable it to move
around silently on the forest oor as it
probes for food. Kiwis are the only birds
to have nostrils at the end of their bills,
and they use these to smell for prey
among vegetation and below the ground.
Because they are
so well adapted to
After sunset the kiwi
living and hunting
hunts for snails, spiders,
on the ground,
and insects. It uses its
kiwis have been
keen sense of smell
described as
and hearing to locate
prey underground.
honorary mammals.

Peregrine falcons have extremely acute

vision and power dive at high speed from
above their prey. Typically a peregrine will
come out of its dive just behind the prey,
aiming to hit the wings of the victim rather
than risk injury with a direct body hit. If the
impact does not kill the bird, it will break
the birds neck with its sharp, hooked bill.
Its diet mainly comprises pigeons, doves,
waterfowl, and gamebirds, as well as
reptiles and small mammals.


A secretary bird stamps on the
ground with its large, stouttoed feet to flush prey
out of hiding.

Falco peregrinus
Apteryx australis lawryi

a Up to 151 2 in (40 cm)

b In forest and coastal scrub, only on Stewart Island off

southern New Zealand.

a Up to 211 2 in (55 cm)

b Worldwide, but absent from New Zealand, high

mountains, deserts, and polar regions.

c Bluish to dark gray, white face with dark stripe.

c Rounded, wingless bird, with long, slender bill and

light brown mottled plumage.

Stamping for prey

Secretary bird
This large bird feeds on snakes such
as adders and even cobras, but will
also consume lizards, amphibians,
rodents, young birds, bird eggs, and
insects. Small animals are eaten directly,
but larger prey is stamped to death before
being consumed. Dangerous prey such as
snakes are rst stunned by being stamped
on, then killed by being pecked behind the
neck. Small creatures are eaten whole,
while larger prey are pinned to the ground
and pulled apart.

long, powerful legs

used for striking and
pursuing prey

Sagittarius serpentarius

a Up to 41 2 ft (1.4 m)
b Open grassland in sub-Saharan Africa.
c Light gray plumage and black flight feathers,

with black feathers at back of head, and long legs.

Surface paddling
White-vented storm-petrel
Despite its small size, this bird is quite a
strong ier, and feeds from the surface of
the ocean, uttering with rapid, almost
batlike, wing beats above the surface of the
water. As it does this, it paddles its yellow
webbed feet up and down on the water
while it looks for planktonic organisms, such
as crustaceans and small shes, to eat. The
white-vented storm petrel lives most of its
life over open water, only coming to land to
breed. It often nests
in hollows or lava
The common name of
tubes but the
this small bird refers to
presence of
the patch of white
introduced rats has
plumage at the base of
threatened its
the tail, that extends
round to the underside.

Oceanites gracilis

a 6 in (15 cm)
b Waters off Chile and Peru, and around the

Galapagos Islands.

c Mainly black with longish, slender dark legs. Nostrils

fused into a single tube on top of the bill.

One of the most distinctive features of

birds of prey is the way they often kill with
their feet. The white-tailed sea eagle feeds
mostly on shes, small mammals, and
other birds, such as ducks. When hunting
for shes, it sits in a good perch and scans
Haliaeetus albicilla

a Up to 35 in (90 cm)
b Coastal areas, but also inland wetlands, rivers, and

lakes, in Europe and Asia.

c Large eagle with brownish body, white tail, and

yellow, hooked bill.

Feet first
Since their diet consists entirely of shes,
ospreys are also known as sh eagles.
They have several special adaptations for
catching shes up to 4 lb (2 kg) in weight.
Their toes are of equal length, and the
outer toe is reversible, allowing the bird to
have two toes hooked into either side of a
shs body. Not only do the feet have sharp
talons to hold slippery shes, they also
have barbed foot pads to increase their
grip. Their hunting technique consists of
ying high above the water to spot shes
below, then, after hovering for a second or
two, diving down to enter the water talonsrst with outstretched legs. Ospreys will
enter water to a depth of more than 3 feet
(nearly a meter) to catch shes.

the water using its excellent eyesight

before swooping down to seize shes
swimming near the surface with its large,
hooked talons. The bottoms of its feet
have sharp outgrowths to help keep a rm
hold of the slippery prey. The eagle takes
food back to its nest or perch, where it
holds it down and rips it apart using its
large, vulturelike bill. Preferring open
coastal locations over inland sites, the
territory of a white-tailed sea eagle may
be more than 50 miles (80 km) across
if prey is not very abundant.
Each adult bird needs to consume
around 18 oz (0.5 kg) of prey each day, but
can survive short lean periods. Although
they are superb hunters, these eagles will

Ospreys can close

their nostrils to
keep water out of
their nasal passages
when diving.
Sometimes ospreys have difficulty
getting airborne from the water.
Once in flight they carry the fish
headfirst to reduce wind resistance.
Pandion haliaetus

a 26 in (65 cm)
b Anywhere there is open water and plenty of fishes,

worldwide except Antarctica.

c Brown upper body and wings, white underside.

sometimes take carrion, searching the

shore for dead sh, or stealing prey from
other predators such as otters and
ospreys. In some areas, these eagles may
compete with golden eagles for rabbits
and hares.
After a spectacular aerial courtship,
which involves the birds cartwheeling
through the air with their talons locked
together, pairs of eagles bond for life and
build a large nest, or eyrie, of twigs and
sticks in a tree or on a sheltered rocky cliff.
Females usually lay one or two eggs, and
the young are fed for three months. The
mortality rate is high with 6070 percent
of the young birds failing to
survive their rst winter.

Swooping low over the water
on its powerful broad wings,
a white-tailed sea eagle can
pluck fish from the surface
without getting itself wet.

long wing tips


Snatching fish
White-tailed sea eagle

This species, also known as the Namib desert sidewinding or

sand adder, hunts by burying itself in the sand with just its tail and
eyeslocated on the top if its headshowing, making good use
of its excellent camouage coloring. The end of the tail is exposed
and this, especially in very dark-tailed examples of the species, is
used as a lure. The preyattracted by the movement and grublike
appearance of the tailis grabbed and injected with venom. This
subdues the prey, which is then swallowed whole. Favorite prey of
this snake include desert lizards and geckoes. As an adaptation
to living on shifting desert dunes, and even to climbing steep
inclines, Peringueys desert adder moves by a process known as
sidewinding. The body is moved in a series of lateral movements,
leaving sinuous marks that are characteristic of the species.
Living in very dry conditions, such as the almost rain-free Namib
desert, the snake gets most of the water it needs to survive from
its prey, particularly lizards, which have a high water content.
However, it also draws a supply
of uid from the condensation
that settles on its body.

Bitis peringueyi

a 1012 in (2530 cm)

f Coastal sands in Angola and Namibia.
c Body pale brown, orange, or sandy, with
gray-brown spots, and paler underside. Eyes
on top of flattened head. Tail typically brown.

Peringueys desert
adder buries itself in
sand (far left and
above) to keep cool
and conserve moisture
as well as to conceal
itself from prey.


A desert lizard has
fallen for the snakes
appetizing-looking tail
lure and is in the
process of being
swallowed whole.

Snakes have rectangular

scales on the underside of
their body, the edges of which
provide traction like tread
on a tire.

Sidewinding is a variation of the
more typical undulating motion
that snakes use to move around,
and is seen especially in species
that live on loose sand in deserts
or slippery mudats where there
are no rm objects such as rocks
against which the snake can push.
By throwing its body into a series
of lateral loops, the snake moves
sideways with most of its body
clear of the surface at any time.
This allows it to move over very
hot sand, leaving behind a series
of J- or S-shaped tracks.


Tail lure
Peringueys desert adder

Group members cooperate when catching and
sharing food and defending their breeding territory
from predators such as ravens, great horned owls
and coyotes. Harriss hawks are not fast fliers but
have excellent vision and hearing.

Sensitive eyes
Common kestrel
The diet of common kestrels consists
mainly of small mammals, such as voles
and mice, but they will also catch small
birds. It might seem that the kestrel would
have a difcult task catching small,
fast-moving prey but they have
evolved a special hunting ability.
The eyes of kestrels are sensitive
to ultraviolet light, and they use
the fact that rodent urine reects
ultraviolet light to nd their prey.
Voles, mice, and other rodents
habitually mark their trails with urine
and feces, so kestrels are

Falco tinnunculus

a 1215 in (3038 cm)

b Moorland, meadow, farmland, and urban areas in

Europe, Asia, and Africa.

c Males are smaller and grayer than the reddish brown

females. Tail has dark bar accentuated by white tip.

Plunge diving
Cape gannet
These birds use a plunge-diving technique
from as high as 98 ft (30 m) above the
surface to catch schooling shes such as
sardines and anchovies. When suitable
shes are spotted, the gannets start a
dive from a height determined by the
depth of the school. Just before they hit
the water, the gannets pull their wings
back and close to the body, to form a
streamlined, arrowhead shape. Having
entered the water, they can chase prey

This hawks usual prey are small mammals

such as ground squirrels and wood rats,
birds, and reptiles, but by hunting together
in groups of up to six individuals, it is able
to take prey as large as jackrabbits, hares,
or large gamebirds. When prey is spotted,
the hawks land and scare the victim from
its hiding place by force of numbers. One
of the group then captures and kills it. A
small number of a hunting group may also
take turns to y ahead to scan for prey.
Group hunting, which